Journal of Hymenoptera research

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Journal of

Hymenoptera
Research

Volume 12, Number 2 October 2003

ISSN #1070-9428
CONTENTS

BENNETT, A. M. R. A new genus and five new species of Neotropical Tryphoninae

(Hymenoptera: Ichneumonidae) 209

BISCHOFF, I., K. FELTGEN, and D. BRECKNER. Foraging strategy and pollen preferences

of Andrena vaga (Panzer) and Colletes cunicularius (L.) (Hymenoptera: Apidae) . . . 220

BRICENO G., R. A. Taxonomic revision of the genus Sesioctonus Viereck (Hymenoptera:

Braconidae: Agathidinae) 238

HARTLEY, C. S. and R. W. MATTHEWS. The effect of body size on male-male combat in

the parasitoid wasp Melittobia digitata Dahms (Hymenoptera: Eulophidae) 272

KULA, R. R. Morphological variation in Opius Wesmael (Hymenoptera: Braconidae) with

an emphasis on nearctic species in the subgenus Gastrosema Fischer 278

PULAWSKI, W. J. Prosopigastra morogoro, a new species from Tanzania (Hymenoptera:

Apoidea: Crabronidae: Larrini) 303

SHARKOV, A., T E. KATZNER, and T. BRAGINA. A new species of Copidosoma

Ratzeburg (Hymenoptera: Encyrtidae) from eagle nests in Kazakhstan 308

SMITH, D. R. and D. H. JANZEN. Food plants and life histories of sawflies of the families
Tenthredinidae and Pergidae (Hymenoptera) in Costa Rica, with descriptions of
four new species 312

SMITH, D. R. and M. S. SAINI. Review of the southeastern Asian sawfly genus Eusunoxa

Enslin (Hymenoptera: Tenthredinidae) 333

VILHELMSEN, L. Larval anatomy of Orussidae (Hymenoptera) 346

NOTE:

AMIET, F and V. MAUSS. First report of male sleeping aggregations in the pollen wasp

Celonites abbreviatus (Villers, 1789) (Hymenoptera: Vespidae: Masarinae) 355

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j. HYM. RES.
Vol. 12(2), 2003, pp. 209-219

A New Genus and Five New Species of Neotropical Tryphoninae

(Hymenoptera: Ichneumonidae)

Andrew M. R. Bennett

Centre for Biodiversity and Conservation Biology, Royal Ontario Museum, 100 Queen's Park,

Toronto, Ontario, Canada, M5S 2C6 [current address: Agriculture and Agri-Food Canada,

K.W. Neatby Building, 960 Carling Avenue, Ottawa, Ontario Canada, K1A 0C6];

email: bennetta@agr.gc.ca

Abstract. — A new genus of tryphonine ichneumonid, Boethella Bennett, n. gen. is described
from the Neotropics. Synapomorphies are provided which support the sister group relationship
of Boethella and Boethus Foerster (Tryphonini) (the latter known from the Holarctic, Neotropical
and Ethiopian regions). Boethella darlingi Bennett, n. sp. is described from eastern Brazil, B.
canilae Bennett, n. sp. (type species) is described from southern Mexico to eastern Brazil, B.
hubleyi Bennett, n. sp. is described from southeastern Brazil, B. guidottiae Bennett, n. sp. is
described from western and central Brazil and B. curriei Bennett, n. sp. is described from Peru.

The latest survey of the family Ichneu-
monidae (Yu and Horstmann 1997) listed
21,805 described extant species classified
into 1485 genera. These genera are as-
signed to 36 or 37 extant subfamilies de-
pending on opinion (Wahl 1990, Yu and
Horstmann 1997, Gauld 2000, Gauld and
Wahl 2000). Some studies have attempted
to elucidate the subfamily relationships
(Wahl 1991, Wahl and Gauld 1998, Quicke
et til. 2000); however, to date, no complete
subfamily phylogeny is available for the
Ichneumonidae. A recent study of one
subfamily, the Tryphoninae, was under-
taken in order to ascertain its placement
within the family (Bennett 2002). In the
process of this study, two new genera
were discovered, one of which is de-
scribed below. The other new genus from
the tribe Oedemopsini will be described
elsewhere allowing a full discussion of the
characters and relationships of the 12 ex-
tant oedemopsine genera.

The subfamily Tryphoninae comprises
1170 described species (Yu and Horst-
mann 1997) assigned to 53 described gen-
era in seven tribes (Bennett 2002). The sub-
family is cosmopolitan with its centre of

diversity in the Holarctic Region (Kaspar-
yan 1973). Tryphonines are koinobiont ec-
toparasitoids of lepidopterans and saw-
flies of the families Xyelidae, Tenthredi-
nidae, Cimbicidae, Diprionidae and Argi-
dae. They exhibit the uniquely derived
trait of bearing eggs that travel down the
outside of the ovipositor (Kasparyan 1973)
(although this trait is unknown in the Idi-
ogrammatini). The apical end of the stalk
(which bears an anchor in most genera)
travels down the inside of the ovipositor
so that the body of the egg is suspended
by the stalk ventral to the ovipositor. This
anchor is pushed through the host integ-
ument during oviposition, thereby hold-
ing the egg in place until hatching. Fe-
males of most genera of tryphonines have
the habit of bearing an egg on their ovi-
positor while searching for their hosts
(Kasparyan 1973) which provides an in-
controvertible character to identify them
as tryphonines. Male tryphonines and fe-
males that do not carry eggs externally
during host searching (e.g. the Phytodie-
tini) are more difficult to recognize be-
cause the subfamily is relatively heter-
ogenous in structure with some taxa re-

210

Journal of Hymenoptera Research

sembling ctenopelmatines, whereas others
resemble phygadeuontine cryptines, ban-
chines or ophionines.

During studies at the American Ento-
mological Institute (AEIC), I found a series
of Neotropical specimens set aside by
Henry Townes labeled "New Genus near
Bocthus". Two of these putative new spe-
cies were included in a cladistic analysis
of tryphonine generic relationships (Ben-
nett 2002) to examine their status and their
hypothesized relatedness to BoetJius. The
cladistic analysis verified the generic sta-
tus of this new genus as well as its sister
group relationship with Boethus (see Ben-
nett 2002, Bennett in prep, and below).
The present paper describes this new tax-
on as Boethella Bennett, n. gen., the 54th
genus of the Tryphoninae (37th genus of
the tribe Tryphonini).

MATERIALS AND METHODS

Specimens were borrowed from and are
deposited in the American Entomological
Institute, Gainesville, FL, USA (AEIC) (D.
Wahl) and the Canadian National Collec-
tion, Ottawa, Canada (CNCI) (J. Huber).
Examination of other major ichneumonid
collections (e.g. INBio Costa Rica and The
Natural History Museum, London) did
not reveal more specimens of this genus.

Morphological terms follow Townes
(1969) with some modifications: supra-an-
tennal area for 'frons', supraclypeal area
for 'face', malar space for 'cheek', epicne-
mial carina for 'prepectal carina' and la-
terotergites for 'epipleura'. MSI, MS2 re-
fers to metasomal segments 1, 2 etc. Tl etc.
refers to the tergites of metasomal seg-
ments 1 etc. and SI etc. refers to the ster-
nites of metasomal segment 1, etc. Wing
venation terms follow the Comstock-
Needham system as updated by Ross
(1936) and incorporates the recommenda-
tions of Goulet and Huber (1993) except
for naming of the vein that forms the dis-
tal edge of fore wing cell l + 2Rs (the 'ar-
eolet' of Townes 1969) which is referred to

as vein 3rs-m in accordance with Gauld
(1997).

TAXONOMIC PLACEMENT

The strongest evidence that a taxon be-
longs to the Tryphoninae is that its fe-
males bear stalked eggs that travel down
the outside of the ovipositor. The egg of
Boethella is not known, therefore the place-
ment of Boetliella in the Tryphoninae is not
certain. Most adult tryphonines exhibit the
following characters: clypeus with an api-
cal fringe of hairs; cell 1 + 2Rs (areolet) of
fore wing triangular to subrectangular;
spiracle of Tl anterior to middle; Tl with
a glymma present; dorsal valve of ovipos-
itor high (not strongly tapered) and un-
notched subapically. Of these characters,
Boethella possesses the clypeal fringe of
hairs (albeit sparse) and the high, un-
notched ovipositor (Fig. 7). The latter
character precludes the placement of Boe-
thella in the Ctenopelmatinae. In addition,
the areolet of the fore wing of Boethella
(Fig. 1) resembles other tryphonines which
have an open areolet (i.e., if the areolet
were closed, it would be triangular, not
pentagonal as in cryptines or ichneumon-
ines or rhombic as in mesochorines). On
the basis of these three characters, Boethella
fits best in the subfamily Tryphoninae
(compared to all other extant subfamilies),
despite its apomorphic structure of Tl
(i.e., glymma absent and spiracle posi-
tioned posteriorly which is the condition
found in cryptines and ichneumonines). It
should be noted that Boethella is not the
only genus of tryphonine with the latter
two character states (e.g., Sphiuetus Grav-
enhorst and Ankylophon Gauld).

Boethella can further be assigned to the
tribe Tryphonini because of its apomorph-
ic pectinate tarsal claws that within the
Tryphoninae are known only in the tribes
Tryphonini, Exenterini (= the Exenterus
group of genera, see Bennett 2002),
Sphinctini and Phytodietini. Boethella does
not possess any apomorphies that would
place it in either of the latter two tribes

Volume 12, Number 2, 2003

211

Fig. 1. Boethella canilae, holotype female, habitus.

(e.g., the strongly pointed apical edge of
the clypeus in the Sphinctini or the loss of
propodeal carinae in the Phytodietini).
Boethella also cannot be assigned to the Ex-
enterini because it has paired tibial spurs
on both the middle and hind legs (the Ex-
enterini have the autapomorphies of only
one spur on the middle leg and no spurs
on the hind leg).

The sister group relationship of Boethella
and Boethus is supported by the following
synapomorphies: occipital carina absent;
epomia absent; notauli absent; fore wing

vein 2m-cu with one bulla; fore wing vein
3rs-m absent (areolet open); Tl petiolate;
spiracle of Tl in posterior 0.4. A complete
cladistic analysis describing character po-
larities and the relationships of all try-
phonine genera including Boethus and Boe-
thella is given in Bennett (2002) and Ben-
nett (in prep.). Boethella can be distin-
guished from Boethus by the former's
possession of propodeal, epicnemial and
submetapleural carinae (all of which are
absent in Boethus). In addition, the glym-
ma is absent in Boethella (present in Boe-

212

Journal of Hymenoptera Research

thus). Boethella is exclusively Neotropical
and the majority of species of Boethus are
known from the Neotropical and southern
Nearctic regions (Townes et ah 1992); how-
ever, several Ethiopian species of Boethus
are known (Scaramozzino 1991) as well as
one Palaearctic species (Kasparyan 1973).

Boethella Bennett, n. gen.

Type species. — Boethella canilae Bennett,
by original designation.

Diagnosis. — Distinguished from other
genera of tryphonines by the combination
of: 1) occipital carina absent; 2) propodeal
carinae present (Fig. 6). In addition, the
ovipositor of Boethella is distinctive within
the Tryphoninae (slightly upcurved with
a high, wide, apically rounded dorsal
valve that strongly overlaps the ventral
valve medially) (Fig. 7).

Description. — Fore wing length 2.9 to 4.8
mm; clypeus slightly rounded in profile,
without a transverse line separating it into
dorsal and ventral faces, apical margin
truncate to slightly rounded in anterior
view, strongly impressed laterally (Figs. 2
and 3) without medial paired tubercles
(medial notch absent), clypeal fringe of
hairs present, but sparse; malar space
obliterated (mandibular socket contiguous
with ventral edge of eye) (Fig. 2) except in
B. darlingi space is 0.5 times basal width
of mandible (Fig. 3); lower mandibular
condyles separated by distance greater
than distance of inner eye margins at level
of clypeal foveae; mandible with teeth
subequal in width and height, moderately
convex in cross-section near base; labio-
maxillary complex moderately elongate,
glossae visible in anterior view in most
specimens (Fig. 1); occipital carina absent;
postgena without a tooth; supra-antennal
area without a horn or carina; antennal
sockets separated by distance greater than
0.5 diameter of socket; eyes without prom-
inent setae; epomia absent (Fig. 1); dorso-
posterior region of pronotum not strongly
thickened in dorsal view; epicnemial ca-
rina present, not dorsally curving toward

anterior edge of mesopleuron (Figs. 1, 4-
5); auxiliary carina of mesopleuron either
long and joining epicnemial carina (Figs.
4-5) or short and not joining (Fig. 1); ster-
naulus present (Figs. 1, 4-5); subtegular
ridge slightly curving out laterally, not
produced into a vertical lamella that near-
ly reaches tegula when tegula is down; no-
tauli absent; projection on posterolateral
edge of mesoscutum absent; carinae of
scutellum present at base only; propodeal
carinae all present except lateral longitu-
dinal carinae absent (Fig. 6), medial lon-
gitudinal carinae strongly raised, medial
portion of posterior transverse carina
weak; submetapleural carina present; fore
tibia without an anterior, apical tooth; fore
tibial spur evenly curved; middle and
hind trochanters two segmented; middle
and hind tibiae each with two spurs; tarsal
claws pectinate to apex or nearly to apex
(Fig. 1); fore wing vein 3rs-m absent (Fig.
1); fore wing vein 2m-cu weakly to strong-
ly inclivous with one bulla (Fig. 1); wings
hyaline to moderately infumate; Tl petio-
late (Fig. 6) with spiracle at 0.6 to 0.75,
dorsal longitudinal carinae absent (Fig. 6),
dorsolateral longitudinal carinae present
but not extending to spiracle (Fig. 1);
glymma of Tl absent (Fig. 1) (slight de-
pression present ventral to dorsolateral
longitudinal carina in some specimens,
but not a glymma); Tl and T2 not fused,
their sculpture impunctate; SI not fused to
Tl, membranous portion of SI not or only
slightly projecting lateral to sclerotized
portion of Tl; T2 without a transverse
postmedian groove or oblique grooves de-
lineating the anterolateral corners; latero-
tergites of MS2 to MS4 separated from ter-
gites by a complete crease; T6 to T8 not
strongly turned anteriorly under T5; ovi-
positor (only known in two species) short-
er than apical depth of metasoma, mod-
erately upcurved, dorsal valve thick and
rounded apically, overlapping ventral
valve laterally (Fig. 7).

Mature larva. — Unknown.

Egg. — Unknown.

Volume 12, Number 2, 2003

213

Hosts. — Unknown. The sister genus Boe- Species included. — Five species (see key
thus has been reared from argid sawflies and descriptions below).
(Townes et al. 1992, Gauld 1997). Etymology. — Boethella is a modification

Distribution. — Southern Mexico, Peru of Boethus (which means "helper" in
and Brazil. Greek) indicating its close relationship

with this genus. Its gender is feminine.

KEY TO THE SPECIES OF BOETHELLA BENNETT

1 Malar space 0.5 times basal width of mandible (Fig. 3) (eastern Brazil)

darlingi Bennett, n. sp.

Malar space obliterated: dorsal edge of socket of mandible contiguous with ventral edge
of eye (Fig. 2) 2

2(1) Mesopleuron with auxiliary carina short, not extending to epicnemial carina (Fig. 1) ... 3
Mesopleuron with auxiliary carina extending from anterior edge and joining epicnemial
carina slightly ventral to ventral edge of pronotum (Figs. 4 and 5) 4

3(2) Hind tibia brown. (Brazil to southern Mexico) canilae Bennett, n. sp.

Hind tibia with basal 0.7 yellow, apical 0.3 brown (Brazil — Santa Catarina)

hubleyi Bennett, n. sp.

4(2) Epicnemial carina extending dorsal to point of union of auxiliary carina by at least the
length of auxiliary carina (Fig. 4). T4 predominantly brown, yellow laterally and with

a yellow medial longitudinal stripe or spot in some specimens (Peru)

curriei Bennett, n. sp.

Epicnemial carina extending dorsal to point of union of the auxiliary carina by much

less than the length of auxiliary carina (Fig. 5), or not extending at all. T4 entirely

yellow or yellow with a trace of brown in apical 0.2 and with a longitudinal brown

or brown and white region in medial 0.3 (Brazil: Mato Grosso and Amazonas) ....

guidottiae Bennett, n. sp.

Boethella darlingi Bennett, n. sp.

Fig. 3

Diagnosis. — Distinguished from other
species of Boethella by having the malar
space 0.5 times basal width of mandible
(Fig. 3) (not zero times basal width).

Female. — Unknown.

Male (holotype). — Fore wing length 3.5
mm; medial part of apical edge of clypeus
slightly convex in anterior view, without
emargination; groove between clypeus
and supraclypeal area weak laterally so
that base of clypeus is relatively flat; malar
space 0.5 times basal width of mandible;
antenna with fourteen flagellomeres; aux-
iliary carina of mesopleuron short, not
joining epicnemial carina; abscissa of fore
wing vein M between 3rs-m and 2m-cu

greater than 0.75 length of 2m-cu; spiracle
of Tl posterior to 0.7. Orange; antenna ex-
cept anterior side of apical two to three
flagellomeres, apical 0.2 of mandible, pos-
terior of occiput medial to inner margin of
eyes, area between ocelli, pronotum along
dorsal edge, lateral lobes of mesoscutum
and anterior 0.3 of medial lobe, scuto-scu-
tellar groove, ventral edge of scutellum,
mesopleuron except dorsoanterior quar-
ter, mesosternum except small region me-
dioposteriorly; dorsal and ventral edges of
metapostnotum, anterior groove of pro-
podeum including base of medial longi-
tudinal carinae, ventroanterior corner of
metapleuron; apical 0.8 of hind tibia, hind
tarsus, wing veins and stigma, T2 except
anterior 0.2, lateral 0.2 and posterior 0.2,

214

Journal of Hymenoptera Research

2. canilae

3. darlingi

Figs. 2-3. Boethella spp. male, anterior view of head.
2, B. canilae, paratype. 3, B. darlingi, holotype.

T3 to T7 except posterior 0.2 brown; clyp-
eus, ventral 0.6 of supraclypeal area, ven-
tral 0.5 of gena, propleuron, ventral 0.2 of
pronotum, fore leg except coxa, entire
middle leg, coxa, trochanter, femur and
basal 0.2 of tibia of hind leg, Tl, anterior
0.2 and lateral 0.2 of T2 yellow; dorsal 0.8
of pronotum except dorsal ridge and me-
dial region of petiolar region of propo-
deum brownish orange; mouthparts ex-
cept apical 0.5 of mandible, anterior side
of apical two to three flagellomeres, fore
coxa and metasomal sternites white; pos-
terior 0.2 of T2 to T6 translucent white;
gonoforceps light brown; wings strongly
infumate basally, fading to hyaline apical-

Material— Holotype 6 : BRAZIL, Rio de
Janeiro State, Guanabara, Rio Grande, Re-
presa, 1-31. iii. 1972 (Alvarenga) (AEIC).

Etymology. — This species is named in
honour of Dr. D. C. Darling, senior curator
at the Royal Ontario Museum, in recog-
nition of his long-standing appreciation of
the magnificence of the family Ichneu-
monidae.

Comments. — Known only from the ho-
lotype. Boethella darlingi may be the sister
species of the other four species of Boe-
thella because it lacks the obliterated malar
space that is synapomorphic of these four
species.

Boethella canilae Bennett n. sp.

Figs. 1-2, 6-7

Diagnosis. — Distinguished from other
species of Boethella by the combination of
all: 1) malar space obliterated (mandibular
socket contiguous with ventral edge of
eye) (Fig. 2); 2) auxiliary carina of meso-
pleuron short, not joining epicnemial ca-
rina (Fig. 1); 3) hind tibia brown.

Female (holotype). — Fore wing length 4.1
mm; medial part of apical edge of clypeus
slightly and broadly emarginate; groove
between clypeus and supraclypeal area
moderately strong laterally; malar space
obliterated (dorsal edge of mandibular
socket contiguous with ventral edge of
eye) (Fig. 2); antenna with sixteen flagel-
lomeres; auxiliary carina of mesopleuron
short, not joining epicnemial carina (Fig.
1); abscissa of fore wing vein M between
3rs-m and 2m-cu less than 0.5 length of
2m-cu; spiracle of Tl anterior to 0.7. Yel-
lowish orange; apical 0.2 of mandible, an-
tenna except apical flagellomere, occiput
posterolaterally, posteriorly and in a lon-
gitudinal stripe extending posteriorly
from between lateral ocelli, dorsal 0.8 of
pronotum, tegula, anterior 0.5 of medial
lobe mesoscutum and all of lateral lobes,
ventral 0.7 of mesopleuron except anterior
to epicnemial carina and in ventroposter-
ior corner, mesosternum except medial
0.3, metanotum, hind tibia and tarsus,

Volume 12, Number 2, 2003

215

4. curriei

5. guidottiae

6. canilae

Figs. 4-6. Boethella spp. 4-5, Male, lateral view of
mesopleuron: aux. = auxiliary carina, epic. = epic-
nemial carina. 4. B. curriei, paratype. 5, B. guidottiae,
paratype male. 6, B. canilae, holotype female, dorso-
posterior view of propodeum and first metasomal
segment.

wing veins and stigma, spot occupying
posterior 0.5 and lateral 0.3 of T2, T3 to T7
except posterior 0.2 and triangular-shaped
medial portions of T4 to T7 brown; head
(except antenna, mouthparts and occiput
as noted above), ventral 0.2 of pronotum,
posterior 0.5 of medial lobe of mesoscu-
tum, scutellum, dorsal 0.3 of mesopleuron,
anterior to epicnemial carina and in ven-
troposterior corner orange; posterior 0.2 of
T2, posterior 0.2 and lateral 0.2 of T3 to
T7, medial triangular portions of T4 to T7,
metasomal epipleura and sterna including

vent.

/

dors./

hyp-

sheath

Fig. 7. Boethella canilae, holotype female, lateral view
of posterior of metasoma showing ovipositor: vent. =
ventral valve of ovipositor, dors. = dorsal valve of
ovipositor, sheath = ovipositor sheath, hyp = hypo-
pygium.

hypopygium whitish yellow; dorsal valve
of ovipositor, ovipositor sheath and mem-
branes at base of ovipositor white; apical
flagellomere light brown; membrane of
wings hyaline with a trace of infumation
apically in fore wing.

Male. — Same as female except fore wing
length 2.9 to 4.8 mm and antenna with fif-
teen to eighteen flagellomeres. Colour var-
iations: Dark morph as female, except all
flagellomeres brown, occiput with less
brown posterolaterally, apical 0.2 of hind
tibia brown in some specimens and T2
completely brown except orange in a tri-
angular region basomedially and yellow-
ish white in posterior 0.2; gonoforceps
light brown. Light morph as female except
clypeus, supraclypeal area and orbits yel-
lowish orange; occiput, pronotum, meso-
scutum, scutellum and mesopleuron com-
pletely orange; T2 entirely brown except
anterolateral corners (or entire anterior
0.1) orange, posterior 0.2 yellowish-white;
gonoforceps light brown. Inter morph as

216

Journal of Hymenoptera Research

light morph except apical flagellomere
only light brown at apex; lateral lobes of
mesoscutum completely brown; dorsal 0.5
of pronotum orange brown.

Material— Holotype 9, BRAZIL, Espir-
ito Santo, Castelo, l-30.xi.1976 (M. Alvar-
enga) (AEIC). Paratypes. 3 6, same data as
holotype; 1 6, Sao Paulo State, Sao Jose
do Barreiro, Serra da Bocaina, 22°37'59"S,
44°34'59"W, 1650 m, l-30.xi.1969, (Alvar-
enga and Seabra) (AEIC); 1 6, MEXICO,
Chiapas, 10 km south of Ocozocoautla,
1220 m, 2.viii.l962. (H. Milliron) (CNCI).

Etymology. — This species is named in
honour of Dr. C. Canil in recognition of
her exemplary volunteer work at the Roy-
al Ontario Museum.

Comments. — Boethella canilae is quite var-
iable in colour with the pronotum, meso-
pleuron and mesoscutum ranging from
predominantly brown to completely or-
ange (in same collection site). The male
specimen from Mexico is smaller than the
other males (fore wing = 2.9 mm) but ap-
pears to be conspecific (similar to the light
morph males except that T2 and T3 are
orange brown instead of brown). The
specimen is in relatively poor condition
and newer material from Mexico may re-
veal structural differences that distinguish
this population as a distinct species.

Boethella guidottiae Bennett, n. sp.

Fig. 5

Diagnosis. — Distinguished from other
species of Boethella, by the combination of:
1) auxiliary carina of mesopleuron long,
joining epicnemial carina, the latter ex-
tending only slightly dorsal to point of
union with auxiliary carina or not extend-
ing (Fig. 5); 2) T4 predominantly yellow.

Female (holotype). — Fore wing length 4.0
mm; medial part of apical edge of clypeus
with a slight, narrow emargination;
groove between clypeus and supraclypeal
area moderately strong laterally; malar
space obliterated (dorsal edge of mandib-
ular socket contiguous with ventral edge
of eye); antenna with seventeen flagello-

meres; auxiliary carina of mesopleuron
long, joining epicnemial carina, the latter
extending only slightly dorsal to point of
union with auxiliary carina (Fig. 5); ab-
scissa of fore wing vein M between 3rs-m
and 2m-cu less than 0.5 length of 2m-cu;
spiracle of Tl anterior to 0.7. Yellow; basal
0.5 of mandible, apical five flagellomeres
of antenna, medial 0.5 of propleuron, me-
dial 0.2 of mesosternum, entire metapleu-
ron, coxa and trochanter of fore and mid-
dle legs, tarsus of fore leg whitish yellow;
posterior 0.2 of T2 to T5 as well as lateral
edges ventral to spiracle, medial triangu-
lar region (widest in posterior) on T3 to
T7, metasomal sternites, hypopygium ex-
cept for a triangular stripe just ventral to
dorsal edge, membranes around oviposi-
tor white; scape, pedicel, apical 0.3 of hind
tibia, hind tarsus, stigma, apical wing
veins and narrow border around medial
triangular regions of T3 to T7 yellowish
brown; supra-antennal area, vertex, occi-
put, dorsal 0.3 of gena, dorsal 0.5 of pro-
notum and mesoscutum yellowish orange;
apical 0.2 of mandibles, flagellum except
apical five flagellomeres, basal wing veins
brown; wing membrane weakly to mod-
erately infumate anteriorly and apically,
hyaline posteriorly and subapically.

Male. — Similar to female except apical
edge of clypeus slightly convex to truncate
medially, (emargination present in only
some male specimens); fore wing length
3.2 to 4.8 mm; antenna with sixteen to
nineteen flagellomeres. Colour variations:
Light morph similar to female except api-
cal two to seven flagellomeres whitish yel-
low and in most specimens, medial 0.5 of
T6 and all of T7 yellowish brown; gono-
forceps whitish yellow. Dark morph or-
ange; clypeus, mouthparts except apical
0.6 of mandible, ventral 0.5 of pronotum,
propleuron, metapleuron, propodeum,
fore leg, middle leg, coxa, trochanter and
femur of hind leg yellow; apical two to
four flagellomeres and sternites of meta-
soma whitish yellow; all structures that
are brown in female are also brown in

Volume 12, Number 2, 2003

217

dark morph male as are all wing veins
and stigma; T2 to T7 vary from yellow to
light brown, tending to be more brown
medially and posteriorly, apical 0.1 to 0.2
of each tergite may be white or uniform
with rest of tergite, some specimens are
also similar to holotype with medial 0.3 to
0.5 of T3 to T7 with unpigmented, trian-
gular regions which may indicate incom-
plete sclerotization of these segments (a
longitudinal, medial suture line is also
present on the posterior tergites in these
specimens); basal 0.7 to 0.8 of hind tibia
may be yellow to brownish yellow (but
base always lighter than apical 0.2 which
is brown); gonoforceps yellowish orange
to whitish brown.

Material— Holotype 9, BRAZIL, Mato
Grosso, Sinop, 12°31'S, 55°37'W, Malaise
Trap, 1-31. x. 1974 (M. Alvarenga) (CNCI).
Paratypes. 5 8 , same data as holotype ex-
cept one from 1-31. x. 1976 and three 1-
31. xi. 1975; 3 8 , same data as holotype ex-
cept 1-31. x. 1976 (AEIC); 2 8 , Amazonas,
4°33'S, 71°38'W, l-30.ix.1979 {Alvarenga)
(AEIC).

Etymology. — This species is named in
honour of Ms. A. Guidotti in recognition
of her dedicated work as technician of the
entomology collection of the Royal Ontar-
io Museum.

Boethella hubleyi Bennett, n. sp.

Diagnosis. — Distinguished from other
species of Boethella by the combination of
rt//: 1) malar space obliterated (mandibular
socket contiguous with ventral edge of
eye); 2) auxiliary carina of mesopleuron
short, not joining epicnemial carina; 3)
hind tibia yellowish orange in basal 0.7,
brown apically.

Female (holotype). — Fore wing length 4.5
mm; medial part of apical edge of clypeus
slightly and broadly emarginate; groove
between clypeus and supraclypeal area
moderately strong laterally; malar space
obliterated (dorsal edge of mandibular
socket contiguous with ventral edge of
eye); antenna with seventeen flagello-

meres; auxiliary carina of mesopleuron
short, not joining epicnemial carina; ab-
scissa of fore wing vein M between 3rs-m
and 2m-cu less than 0.5 length of 2m-cu;
spiracle of Tl anterior to 0.7. Yellowish or-
ange; apical 0.2 of mandible, antennae ex-
cept apical two flagellomeres, anterior 0.5
of medial lobe of mesoscutum, lateral lobe
of mesoscutum except anterior, lateral and
medial edges, mesopleuron, mesosternum
except medial 0.2, apical 0.3 of hind tibia,
hind tarsomeres, wing veins, stigma and
T3 to T7 except posterior 0.2 and lateral
0.1 to 0.2 brown; occiput medioposterior-
ly, pronotum, tegula, posterior 0.5 of me-
dial lobe of mesoscutum, anterior, lateral
and medial edges of lateral lobes of me-
soscutum, scutellum, T2, lateral 0.2 of T3,
lateral 0.1 of T4 to T7 orange; palpi, basal
0.8 of mandibles, fore and middle coxa
and trochanter and anterior 0.7 of Tl whit-
ish yellow; apical two flagellomeres light
brown; membrane of wings hyaline with
a trace of inhumation apically in fore wing.

Male. — Unknown.

Material— Holotype 9, BRAZIL, Santa
Catarina, 27°11'S, 52°23'W, 300-500m,
25.viii.1962 (F. Plaumann) (CNCI).

Etymology. — This species is named in
honour of Mr. B. Hubley in recognition of
his tireless work as collection manager of
entomology at the Royal Ontario Muse-
um.

Comments. — Known only from the ho-
lotype. Colours in fresh material may be
more contrasting because of the age and
condition of the holotype at time of de-
scription. Boethella hubleyi is similar to B.
canilae but the former can be distinguished
by the bi-coloured hind tibia. Similar col-
ouration of the hind tibia is only known
in B. guidottiae, however this can be distin-
guished from B. hubleyi by the structure of
the epicnemial carina.

Boethella curriei Bennett, n. sp.

Fig. 4

Diagnosis. — Distinguished from other
species of Boethella by the combination of:

218 Journal of Hymenoptera Research

1) the auxiliary carina of mesopleuron ange are more yellowish orange and yel-
long, joining epicnemial carina, the latter lowish orange structures more yellow;
extending dorsal to point of union of aux- hind tibia yellow in basal 0.2, yellowish
iliary carina by at least the length of aux- brown medially, brown apically; medial
iliary carina (Fig. 4); 2) T4 predominantly longitudinal stripe on T4 to T7 less prom-
brown, inent; T4 completely yellowish brown ex-

Female. — Unknown. cept for anterolateral corners and posteri-

Male. — Fore wing length 3.2 to 3.8 mm; or 0.2.
medial part of apical edge of clypeus Material. — Holotype 6, PERU, Cusco,
slightly and broadly emarginate; groove near Marcapata, Avispas, 1-30. ix. 1962 (L.
between clypeus and supraclypeal area Vena) (AEIC). Paratypes. 1 6, same data
moderately strong laterally; malar space as holotype except 1-15. x. 1962; 1 6 , same
obliterated (dorsal edge of mandibular as holotype except 20-30. ix.1962.
socket contiguous with ventral edge of Etymology. — This species is named in
eye); antenna with sixteen flagellomeres; honour of Dr. D. C. Currie, curator and
auxiliary carina of mesopleuron long, join- keeper of black flies at the Royal Ontario
ing epicnemial carina, the latter extending Museum, in recognition of his unfathom-
dorsal to point of union of auxiliary carina able appreciation of fried spam sandwich-
by at least the length of auxiliary carina; es.

abscissa of fore wing vein M between 3rs- Comments. — The specimen caught 1-

m and 2m-cu less than 0.5 length of 2m- 15.X.1962 is not only lighter in colour than

cu; spiracle of Tl anterior to 0.7. Yellowish the other two specimens, but also has a

orange; apical 0.2 of mandible, antenna ex- much less pronounced sternaulus and a

cept for apical one to two flagellomeres, weaker, shorter epicnemial carina. Addi-

hind tibia (except in basal 0.2 in some tional material is necessary to determine if

specimens), hind tarsus, wing veins and these differences are intra- or interspecific,
stigma brown (ventral part of stigma light

brown in some specimens); dorsal 0.5 of ACKNOWLEDGMENTS

head, dorsal 0.5 of pronotum and meso- Funding for this study was provided by an NSERC

SCUtum orange (head and pronotum blend operating grant to Dr. D.C. Darling and by the amaz-

uniformly from orange dorsally to yellow- ing and orten under-appreciated efforts of my gain-

ish orange ventrally); basal 0.8 of mandi- fully emPloyed wife Dr. C. Canil. Additional funding

ii „ ^„i • c a ■jji l was provided bv the University of Toronto, Depart-

bles, palpi, fore and middle legs, propo- , v c~ , „ , * „ . ' /

, i i n ^ <- rnn t-, & ' r r ment of Zoology as well as by the Board of the Amer-

deum, lateral 0.2 of T2 to T4, posterior 0.1 ican Entomological Institute which funded travel to

to 0.2 of T3 to T4 and thin, medial longi- the AEIC. The hospitality of Dr. D. Wahl is much

tudinal Stripe on T4 to T7 yellow; tegula, appreciated during visits to the AEIC as well as his

medial 0.6 of T4, (except for posterior 0 1 permission to include undescribed Townes material

to 0.2 and medial stripe), medial 0.8 of T5 !n ,my, studies.In addition, I*. J. Huber was of great

, . , ... F , ' „ , rri, , m help during visits to the CNCI and Dr. N. Johnson

(except for medial stripe), all of T6 and T7 helped pinpoint Bra2llian locallties Two anonymous

(except for medial Stripe) yellowish reviewers made valuable comments to the manu-

brown; apical one to two flagellomeres script. Study space and equipment was provided by

and gonoforceps whitish brown; sternites the R°yal °ntario Museum.
of metasoma yellowish white; glossa
white; wings strongly infumate basally
and dorsally, hyaline subapically and pos- Bennett, A. M. R. 2002. Cladistics of the Tryphoninae
teriorly and weakly infumate apically. (Hymenoptera: Ichneumonidae) with a discus-
Colour variation: specimen caught 1- sion of host use and the evolution of parasitism

iCwir>£'-> ■ i- Ui. . in the Ichneumonidae, Ph. D thesis. University of

lb.x.1962 is lighter than other two speci- Toronto/ 366 pp

mens— structures described above as or- Bennett, A. M. R. in prep. Cladistics of the Tryphon-

LITERATURE CITED

Volume 12, Number 2, 2003

219

inae (Hymenoptera: Ichneumonidae). Memoirs of
the American Entomological Institute.
Gauld, I. D. 1997. The Ichneumonidae of Costa Rica,

2. Memoirs of the American Entomological Institute,
57: 1-485.

Gauld, 1. D. 2000. The Ichneumonidae of Costa Rica,

3. Memoirs of the American Entomological Institute,
63: 1-453.

Gauld, I. D. and D. B. Wahl. 2000. The Townesioni-
nae: a distinct subfamily of Ichneumonidae (Hy-
menoptera) or a clade of the Banchinae? Trans-
actions of the American Entomological Society, 126:
279-292^

Goulet, H and J. T. Huber. 1993. Hymenoptera of the
World: An Identification Guide to Families. Research
Branch of Agriculture Canada Publication 1894/
E, 668 pp.

Kasparyan, D. R. 1973. Fauna of the USSR Hymenop-
tera Vol. Ill Number 1. Ichneumonidae (Subfam-
ily Tryphoninae) Tribe Tryphonini. Nauka Pub-
lishers, Leningrad (In Russian). Translated into
English (1981) by Amerind Publishing Co, Ltd.
New Delhi, 414 pp.

Quicke, D. L. J., M. G. Fitton, D. G. Notton, G. R.
Broad, and K. Dolphin. 2000. Phylogeny of the
subfamilies of Ichneumonidae (Hymenoptera): a
simultaneous molecular and morphological anal-
ysis. In: Austin, A. D. & Dowton, M. (eds) Hy-
menoptera: Evolution, Biodiversity and Biological
Control. CSIRO, Collingwood, Victoria, pp. 74-
83.

Ross, H. H. 1936. The ancestry and wing venation of
the Hymenoptera. Annals of the Entomological So-
ciety of America, 29: 99-111.

Scaramozzino, P. L. 1991. Two new species of the ge-
nus Boethus Foerster, 1869 from Africa (Hyme-
noptera, Ichneumonidae, Tryphoninae). Bollettino
delta Soeieta Entomologica Italiana. 123: 55-61.

Townes, H. K. 1969. Genera of Ichneumonidae Part I.
Memoirs of the American Entomological Institute, 11:
1-300.

Townes, H. K., V. K. Gupta, and M. J. Townes. 1992.
Nearctic Tryphoninae. Memoirs of the American
Entomological Institute, 50: 1-296.

Wahl, D. B. 1990. A review of the mature larvae of
Diplazontinae, with notes on larvae of Acaeniti-
nae and Orthocentrinae and proposal of two new
subfamilies (Insecta: Hymenoptera, Ichneumon-
idae). Journal of Natural History, 24: 27-52.

Wahl, D. B. 1991. The status of Rhimphoctona with
special reference to the higher categories within
Campopleginae and the relationships of the sub-
family (Hymenoptera: Ichneumondae). Transac-
tions of the American Entomological Society, 117:
193-213.

Wahl, D. B. and I. D. Gauld. 1998. The cladistics and
higher classification of the Pimpliformes (Hy-
menoptera: Ichneumonidae). Systematic Entomol-
ogy, 23: 265-298.

Yu, D. and K. Horstmann. 1997. Catalogue of world
Ichneumonidae (Hymenoptera). Memoirs of the
American Entomological Institute, 58: 1-1558.

J. HYM. RES.
Vol. 12(2), 2003, pp. 220-237

Foraging Strategy and Pollen Preferences of Andrena vaga (Panzer)
and Colletes ciinicularius (L.) (Hymenoptera: Apidae)

Inge Bischoff, Kerstin Feltgen, and Doris Breckner

(IB) Zoologisches Forschungsinstitut und Museum Alexander Koenig, Adenauerallee 160,

D-53113 Bonn, Germany, email: i.bischoff.zfmk@uni-bomi.de;

(KF) Dottendorferstr. 29, D-53123 Bonn, Germany, email: kerstin.feltgen@web.de;

(DB) Johann-Sebastian-Bach-Str. 4, D-77654 Offenbach, Germany

Abstract.— Andrena vaga (Panzer) and Colletes ciinicularius (L.), both vernal ground nesting bees,
were studied in the years of 1996-1999 in a lowbush heath near Cologne, western Germany. Both
species are solitarily but nest gregariously and sometimes form large aggregations with thousands
of nests. They are reported to feed strictly oligolectic on the genus Salix. We observed the daily
foraging rhythms of both species and compared their foraging strategies. Colletes ciinicularius start-
ed provisioning trips earlier in the morning, made more trips per day than A. vaga, and finished
nest provisioning later in the evening. Colletes ciinicularius burrowed even in the dark after 08.00
p.m. Andrena vaga collected pollen and nectar on different days each. One pollen day included 1
to 5 pollen trips. There was no clear correlation between the number of pollen trips and the
occurrence of a subsequent nectar day. We found also no correlation between the occurrence of
nectar-provisioning trips and weather conditions. Pollen loads of both species were analyzed
qualitatively and quantitatively with a cell counter and two different hand-counting systems.
Andrena vaga collected nearly twice as much pollen as C. cuuiciilarius during one foraging trip.
Cells and pollen loads of C. ciinicularius contained large portions of other pollen types, mostly
Rosaceae such as Prunus, Sorbus and Pyrus or Acer, Quercus and Ilex. Thus, C. ciinicularius is not
oligolectic as described in the literature. The percentage of pollen types other than Salix increased
at the end of the flowering period of Salix, which indicates a resource restriction at the end of the
season. The reproductive success of C. ciinicularius measured by nest provisionment exceeded that
of A. vaga, because of longer activity per day and digging activity in the evening.

Andrena vaga (Panzer 1799) and Colletes
ciinicularius (L.) are univoltine, vernal sol-
itary bees which are distributed through-
out the entire Paleartic Region. They pre-
fer to nest in sandy soils and often form
large aggregations (Friese 1923, Moeschler
1938, Vleugel 1947, Westrich 1990). Both
species are reported to be specialized on
Salix as a pollen resource for their larvae
(Westrich and Schmidt 1987). Being spe-
cialized on the same host-plant, their sea-
sonal activity strongly overlaps (Westrich
1990). Therefore, we investigated the di-
urnal activity pattern and foraging strate-
gy of both species. Several studies have
dealt with diurnal activity patterns and
the impact of weather conditions on a

number of mostly Nearctic bee species (cf.
Batra 1999, Lind 1968, Michener and Ret-
tenmeyer 1956, Schonitzer and Klinksik
1990, Stephen 1966), but precise data on
the life cycle of European species of An-
drena and Colletes are very scarce (Geb-
hardt and Rohr 1987, Malyshev 1927, Witt
1992).

By analyzing thoroughly the daily activ-
ity patterns, niche differences between two
species may be shown. Levermann et al.
(2000) investigated the diurnal cycle and
niche differentiation of Dasypoda hirtipes
(Fabricius) and Panurgus calcaratus (Scopo-
li). They demonstrated that apart from
weather conditions, body mass and pollen-
collecting apparatus are important factors

METHODS

Volume 12, Number 2, 2003 221

determining the diurnal activity cycle. Re- and overlaying quicksand, soils are most-
garding solitary Hymenoptera, female size ly sandy, permeable, and poor in nutri-
appears to have a great influence on pro- ents. The climate is humid-oceanic with an
visioning- and reproductive success (Al- annual mean temperature of 9.5 °C and an
cock 1979). Colletes cunicularius is larger annual mean precipitation of 804 mm.
than A. vaga and has more hairs, especially Due to drainage and loss of traditional ag-
on the thorax. Therefore, we hypothesized ricultural use, grass and bushland domi-
that better thermoregulatory abilities allow nate great areas of the heathland (Inter-
C. cunicularius an earlier start to provision- kommunaler Arbeitskreis Wahner Heide
ing activity in the morning. 1989). The investigated aggregations of C.

Apart from the different sizes of the two cunicularius and A. vaga lie on sandy in-

species, the specific pollen-collecting ap- land dunes,
paratus suggests differences in their pol-
len collecting efficiency. Although both
species have trochanter-femur-baskets, Field work. — To study their diurnal ac-

floccus and thoracic pollen basket of A. tivity, individual female bees were

vaga is more strongly developed than in marked with opalith-plates. A total of 238

C. ciniicularius. In a number of bee genera, females of C. cunicularius in 1996 and 174

the size and the type of pollen-collecting females of A. vaga in 1997 were marked,

apparatus results in different amounts of Corresponding nests were identified by a

pollen transported (Braue 1916, Friese colored nail of the same number as on the

1923, Michener et al. 1978, Pasteels and opalith-plate of the female. To record the

Pasteels 1979, Westerkamp 1987, 1996, exact departure and return times of the fe-

Westrich 1990). Thus, we examined the males, several nests were covered with

number of collected pollen grains in both preserving jars (cf. Witt 1992).
species. Climatic parameters were measured

Although both species are regarded as with data loggers (Orion Tiny Logger
strictly oligolectic on Salix (Vleugel 1947, Manager OTLM Tiny talk©). Soil temper-
Westrich and Schmidt 1987, Westrich atures of all years were recorded 20 cm
1990), anecdotal observations reviewed by below the surface, and air temperature
Mader (1999) suggest that C. cunicularius and atmospheric humidity in a portable
visits also flowers other than Salix. A clear weather station were measured at a height
evaluation about oligolecty in a bee spe- of 2 m in 1997-1999. Data from the Co-
des can only be made by a quantitative logne/Bonn airport weather station were
analysis of pollen loads or cell provisions, also used. The data on the activity and the

The aims of this study were to analyze nectar trip frequency of the bees were test-
niche differentiation of two synchronous ed for possible correlation against various
and syntopic bee species on the basis of climatic parameters (daily hours of sun-
activity patterns and the use of host shine, mean daily temperature, mean dai-
plants, to asses the degree of oligolecty, ly atmospheric humidity, humidity of soil
and to compare their pollen collecting ef- and minimal daily soil temperature),
ficiency. Since the activity of most of the bees is

The study area is part of the so-called extremely influenced by weather condi-

"Wahner Heide", a large 5000 ha heath- tions (Larsson 1991, Lind 1968) we created

land, east of the river Rhine near Cologne, a measure for the bees activity indepen-

Since 1961 the heathland has been a mili- dent from weather conditions (cf. Midler

tary training area for NATO and was des- 1994), which we referred to a so-called

ignated a Nature Reserve in 1968. Due to "bee day" (= BD). Such a measure makes

the presence of Quaternary gravel, sand it also easier to compare different years.

222

= bee in the nest

C. cunicularius

n10 n423 n465

first departure 8:35
(n35)

A.vaga
("nectar-day")

A.vaga
("pollen-day")

= provisioning trip

Journal of Hymenoptera Research
waiting at the entrance yyfa = digging

l l l I I I

last return 17:31
(n.20)

n19 n39

tirst departure 11:13
(n25)

last return 14:29
(n39)

n19 n90

n82

H 1 1 h

first departure 10:18
(n20)

H 1 1 1 1 h

H 1 1 1 1 1 1 1 h

last return 16:40
(n29)

H 1 1 1 h

H 1 1 1 1

Time (MET)

Fig. 1. "Ideal ethogram". Time schedule of provisioning behavior of A. vaga and C. cunicularius (dotted line
= end of burrowing unknown). Further information's see text.

One bee day corresponds to a day with
optimal weather conditions, which fe-
males could use completely for provision-
ing activities. A bee day for C. cunicularius
had between 8-9 hours. The measure for
these 8-9 hours was the observed flying
activity at such an optimal day. Other
days with less optimal flying conditions
were defined as follows:

1.0 BD = flying activity of 8-9 hours
0.75 BD = flying activity more than Vi

of one BD
0.25 BD = flying activity less than Vi of

one BD
0 BD = no flying conditions the

whole day

The classification of the bee day of A.
vaga was more difficult because of the
greater variance in activity time, in spite
of good weather conditions:

1.0 BD = flying activity of 8-11 hours

0.75 BD = flying activity more than Vi

of one BD
0.25 BD = flying activity less than Vi of

one BD
0 BD = no flying conditions the

whole day

Nectar- and pollen days of A. vaga. — To
assess a pattern between A. vaga's nectar-
and pollen-provisioning trips, we ana-
lyzed the number of pollen days as well
as the number of pollen trips between nec-
tar trips. One nectar trip corresponds to
one nectar day and the patterns of pollen
trips between nectar trips occurs over a
period days since pollen and nectar are
collected on different days. We obtained
complete observations of pollen trips be-
tween nectar trips of 7 different females.

Pollen analysis. — For pollen analysis we
excavated 10 cells of C. cunicularius in 1996
and 6 cells in 1998. The cells could not be
attributed to a specific nest or time of com-

Volume 12, Number 2, 2003

223

pletion. Additionally, females were cap-
tured to analyze pollen loads. This was
necessary for A. vaga because cells could
not be excavated without destruction and
partial loss of the pollen mass. A total of
38 pollen loads of A. vaga and 28 pollen
loads of C. cunicularius were analyzed (see
Table 1). To remove the complete pollen
load from the bees, all body parts (legs
and sometimes thorax without wings)
were sonicated in vials filled with a liquid
medium (cf. Buchmann and Shipman
1990). For a comparison of grain numbers,
only pollen loads with nearly 100% Salix
were used because the number of pollen
grains varies with their size (see also Sil-
veira 1991, Tasei 1973). For 10 pollen sam-
ples of C. cunicularius also volumetric per-
centages of the different pollen types were
considered (cf. Buchmann and O'Rourke
1991). Pollen grain dimensions of Salix,

Quercus robur and Prunus padus were mea-
sured for 10 grains of each species under
a scanning electron microscope (SEM).
The average size of the grains of Acer
pseudo-platanus were taken from Cromp-
ton and Wojtas (1993).

The number of pollen grains in cells and
pollen loads was counted by different
methods:

a) Ratio-counting with Lycopodium
spores: For this quantification, pollen
was acetolyzed (Erdtmann 1960,
Moore et al. 1991). During acetolysis
a tablet with a known number of Ly-
copodium spores was added (Stock-
marr 1971). In a subsample all pollen
grains and spores were counted on a
slide under a microscope. The total
number of pollen grains in the sample
was estimated from the equation:

total number of grains

added Lycopodium spores
counted Lycopodium spores

X counted grains

b) Cell counter: Most of the samples
were additionally analyzed using a cell
counter (Casy® 1 Cell counter and An-
alyser) and then checked with the SEM.
Eight electronically counted samples for
the years 1996 and 1998 were checked
with the Lycopodium spores method de-
scribed above. Between 500 and 1000
Salix grains were counted, respectively.

c) Hand-counting with a counting
chamber: Samples from 1999 were
counted with a Buerker counting
chamber, a special slide with a cavity
of a defined volume. To achieve an ac-
curacy of 10 or 20 grains per |jl1, we de-
termined the number of subsamples
needed using the following formula (/?
= random sample, s = standard error,
d = accuracy):

//

1.96 X s

d

In this study, 6 to 14 subsamples had to
be counted.

To calculate the number of provisioning
trips for one cell, the average weight of
food stored in the cell has to be divided
by the average weight of the pollen load
carried by the female (cf. Mohamed 1973).
Therefore we determined the dry weight
of the pollen load samples of the year
1996. The females were dried in a drying
chamber, head and wings removed, and
the rest weighted. Then pollen was re-
moved from the body hairs and scopae
with a sonicator. After removing the pol-
len, the thorax and abdomen were dried
and weighed again. The difference corre-
sponds to the weight of the pollen load.

Statistics. — Mean values of all departures
and returns or other activities were used to
construct an "ideal ethogram". Except in
cases when data were not normally distrib-
uted, the median was used. The compari-

224

Journal of Hymenoptera Research

26-
24-

22-

20-

18-

16"
14-

12-

10-

8-

u 6-

? . 4-
0) 2-

3 0

Andrena vaga

First departure

Last return

-i r

i i

07:00 08:00 09:00 10:00 11:00 12:00 13!00 14:00 15:00 16:00 17:00 18:00 19:00 20:00

0)

a

E 26 n

H 24-

22-

20-

18"

16-
14-

12-
10-
8-
6-
4-
2-

Colletes cunicularius

First departure

♦ • 2m
O O -20 cm

0

07:00 08:00 09:00 10:00 1:00 12:00 13:00

— i 1 1 1 i

14:00 15:00 16:00 17:00 18:00

— i 1

19:00 20:00

Time [MET]

Fig. 2. Correlation of time of first departure and last return of A. vaga and C. cunicularius with temperature
at 2 m height (BC = black circle; BS = black square) and 20 cm below soil surface (WC = white circle; WS
= white square). (First departure: BS y = 26.49x + 0.21, r = 0.48, n = 36; WS y = 30.92x-6.38, r = 0.82, n =
38; BC y = - 21. 57x+ 21 .45, r = 0.36, n = 78; WC y = -24.56x+22.37, r = 0.71, n = 78; Last return: BS y =
29.94x-4.31, r = 0.68, n = 42; WS y = 29.35x-6.16, r = 0.81, n = 42; BC y = 0.34x + 19.25, r = 0.01, n = 20;
WC y = 30.12x-3.88, r = 0.89, n = 20).

son of mean values of activity data was
presented in boxplots. These independent
samples were analyzed with the t-test.
Data of pollen samples were treated like-
wise. Mean values of non normally distrib-
uted data (number of pollen grains per pol-
len load and soil temperatures) were com-
pared using the Man-Whitney U-test. The

relationship between activity data and cli-
mate and mixed pollen cells and the
blooming time of Salix were tested by Pear-
sons's correlation analysis.

RESULTS

Foraging strategy — diurnal cycle. — A
comparison of the "ideal ethogram" (Fig.

Volume 12, Number 2, 2003

225

18.00
17:00
16.00
15:00
14:00
lu 13.00
— 12:00

CD

.1 11:00
*" 10:00

09:00
08:00

First departure

Last return

C. cunicularius
A. vaga (pollen)
A. vaga (nectar)

Provisioning flights

Daily flight activity

Fig. 3. Variance and difference in time of first departure, last return, time of provisioning trips and hours of
daily activity of A. vaga and C. cunicularius.

1) shows a different diurnal cycle of A.
vaga and C. cunicularius. For A. vaga two
separate cycles are shown since it collects
pollen and nectar on different days and
this is represented by different flight pat-
terns.

Colletes cunicularius started its first trip
at 8:37 in the morning, after waiting about
20 minutes at the entrance. It made seven
provisioning trips and returned from its
last trip at 17:34. It did not interrupt its
foraging cycle by digging or other activi-
ties in the nest, as indicated by short du-
rations of presence in the nest (six minutes

on average). In the evening, after the last
return to the nest, many females began
burrowing and continued even in the dark
(Fig. 1). Colletes cunicularius showed lower
variation in the number of trips per day
than A. vaga. The first departure and the
last return of C. cunicularius correlated
with the soil temperature (Fig. 2). The fe-
males started earlier when soil tempera-
tures were higher and returned later from
their last trip when temperature was still
high. The correlation of the last return
with the air temperature was not signifi-
cant. The soil temperature at the aggre-

Cells
C.c.

Pollen loads

Year

C.c.

A.v.

1996

10

9

7

1998

6

1

14

1999

18

17

22(_ Journal of Hymenoptera Research

Table 1. Number of excavated cells and collected for the first trip of A. VClga was 8 °C, the
pollen loads (C. c. = C. cunkularius, A. v. = A. vaga). mean temperature of first departure was

12 °C.

To summarize, the females of C. cuni-
cularius started their provisioning behav-
ior earlier in the morning, made more but
shorter trips a day, remained for shorter
periods in the nest, returned later in the
evening (Fig. 3) and burrowed after dark,
gation of C. cunkularius was significantly Andrena vaga started its first trip later in
higher (U = 88698, p < 0.001) than the soil the morning but the temperature thresh-
temperature of the location of A. vaga's ag- old for the first trip was lower than that
gregation, though the year 1996 was much of C. cunkularius.

colder than 1997 (Bischoff 2000). Thus, Nectar and pollen days of A. vaga.— A.
lowest soil temperature during first de- vaga made 1 to 9 pollen trips between two
parture of C. cunkularius (11 °C) differed nectar trips (mean: 4 trips, Table 2). Two
highly from soil temperature in the aggre- groups of flight patterns seem to exist: the
gation of A. vaga (5 °C). The temperature nrst group of bees makes 1 to 4 pollen
threshold for the first trip of C. cunkularius trjpS between two nectar trips, and the
was 9.5 °C; the mean temperature of first second group makes 7 to 9 pollen trips be-
departure was 12 °C. tween each nectar trip.

On pollen days A. vaga made three trips The activities 0f A mga females at dif-
on average. After remaining a while at the ferent bee days are shown in Rg 4 There
entrance, it started its first trip at 10:32. wag fl Wgh percentage of mactive females
The females stayed on average half an qr Q25 bd% [e dayg ^ legs ^ 5 5

our ^ ' , ,. . hours of good flying conditions. On days

two provisioning trips and no digging ac- .Ll_ f a ,.t. ,., n n^,

. . r , , ■ i ■ , with good flying conditions (1.0 BD), more

tivitv was observed during these periods. , , , ,, , . ,i

.,/,.„., , , , ., , , , . j females made pollen trips than nectar

At 16:17 it came back from its last trip and . . , , . , . , c ,

, j . ^ j trips. Yet, this relation was also found on

closed its nest entrance. On nectar days, , . ,

females emerged not before 11:30 and re- °'25 BD s" Furthermore we compared the
turned at 14:29. In the evening no intense Percentage of nectar trips with the bee day
digging activity as observed for C. cum- status of the preceding day. After a 0 BD
cularius, occurred. The last return of the <no flymg conditions the whole day), fe-
females was influenced by the tempera- males made significantly more often a
ture; a significant correlation of the time nectar triP (r = -0.471, p = 0.031, n = 21).
of the last return both with soil tempera- we also tried to correlate the percentage
ture and air temperature was found. The of nectar trips of A. vaga with climatic pa-
first flight in the morning, which started rameters of the same day and the preced-
much later than that of C. cunkularius, ing day (Table 3). At days with more
seems not to be influenced by the temper- hours of sunshine females made more of-
ature at all since a positive correlation was ten a nectar trip but the correlation was
found (Fig. 2). The temperature threshold not very strong. Otherwise no significant

Table 2. Number of observed pollen trips between two nectar trips of A. vaga.

Pollen trips 12345 6 789

Frequency 232100221

Volume 12, Number 2, 2003

227

Table 3. Correlation (Pearson) of the percentage of nectar trips (of all active females of A. vaga) with climatic
parameters of the day of the nectar trip (a) and of the preceding day (b) (n = 20).

Hours of
sunshine

Mean
temperature

Mean humidity

Minimum
temperature

Day
r

P

a
-0.578
**0.008

b

-0.169
0.476

a

-0.86
0.719

b
0.76
0.751

a
0.308
0.186

b
0.244
0.300

a
0.302
0.196

b
0.113
0.636

correlations with climatic parameters were
found.

Composition of pollen loads. — In 1996, 40%
of all analyzed cells of C. cunicularius con-
tained more than 20% of pollen types oth-
er than Salix. Since only percentages of
foreign pollen types lower than 5% (Wes-
trich 1990) are considered as contamina-
tion, we decided to analyze more cells of
C. cunicularius and pollen loads of A. vaga.
In subsequent years, high percentages of
non-Salix pollen were identified in the
cells of C. cunicularius. In fact three of six
excavated cells in 1998 and one of 10 cells
in 1996 respectively, contained no Salix
pollen at all. Only three cells of both years
were pure (>90% Salix) and 9 cells were
mixtures of Salix and other grain types.
The remaining pollen types in the cells of
1996 were mainly composed of various
Rosaceae (Table 4). Apart from Rosaceae,
only Quercus and Sambucus occurred in
higher percentages. Ilex pollen dominated
one pollen load sample of C. cunicularius
of 1996.

Percentages of other pollen types were
also found in pollen loads of C. cunicular-
ius of the years 1998 and 1999. The loads
of two females (captured on 29.4. and
7.5.1999) contained only grains of Acer sp.,
one load (also from 7.5.1999) contained
Acer and Ilex grains in a ratio of 1:1, and
two pollen loads contained exclusively
Quercus pollen. Regarding also the vol-
ume of the different pollen types (10 sam-
ples of 1999), Salix grains represent even a
smaller proportion of the diet of C. cuni-
cularius, since Acer pseudo-platanus, Primus
padus and Quercus robur grains are much
larger than Salix (Fig. 5). Percentages of

other pollen types in pollen loads of C.
cunicularius (in the year 1998) increased
significantly with time, i.e. with the end of
the blooming of Salix (r = 0.731, p = 0.01,
n = 14).

In the graphs of the cell counter (Fig. 6),
mixed loads of mainly two pollen types
{Salix and various Rosaceae) could be rec-
ognized as two separated peaks. These
graphs are counts from C. cunicularius
cells of 1999. They show three clearly sep-
arated peaks. The first peak represents
particles smaller than 10 (xm and can be
interpreted as pollution. The second peak
(15-22 fxm) represents the Salix grains. The
third peak (25-35 |x,m) shows bigger
grains, e.g. Rosaceae. The broad distribu-
tion of grain sizes as displayed in the
counter graph corresponded to different
pollen types detected under the light mi-
croscope. However, results of the hand
countings differed often from percentages
given by the counter. All results of the
counter had to be checked at least quali-
tatively by microscope, because one peak
could represent pollen types other than
Salix.

The cell counter calculated a mean num-
ber of 1512901 (± 720715 SD) pollen grains
per pollen load for C. cunicularius. Only 7
out of 18 pollen loads counted by the cell
counter were pure Salix loads. In A. vaga,
electronic counting determined 2058692
(± 737197 SD) grains per pollen load. An-
il rena vaga collected on average one and a
half more pollen on one provisioning trip
than C. cunicularius. This difference is
highly significant (t-test, p < 0.001).

A comparison of the results using dif-
ferent counting methods is displayed in

228

Journal of Hymenoptera Research

0,25 BD □ 0,75 BD

1.00BD

77

Nectar trip

Pollen trip

Resident (in nest)

Fig. 4. Activities of A. vaga females at a bee day of the category 0,25 (< 5,5 hours good flying-conditions),
a bee day of the category 0,75 (> 5,5 hours good flying conditions) and the category 1 (8-11 hours good
flying-conditions) (number of total observations = 360).

Table 5. Ratio counting with Lycopodium
spores showed greater divergence from
cell counter results than counting with the
counting chamber. Counting chamber re-
sults revealed a mean difference of 255728
(± 192194) grains in comparison to the cell
counter. Regarding Lycopodiiun ratio
counting, one half of the results exceeded
the cell counter calculations and the other
half was below the cell counter calcula-
tions. The mean difference between Lyco-
podium spores ratio counting and cell
counter was 498713 (± 675670) grains.

The pollen loads of A. vaga contained
fewer foreign grains than those of C. cun-
icularius. In five (13%) of 38 loads, we
found percentages of other pollen types

ranging from 1 to 7% and consisting most-
ly of Rosaceae, Quercus and Betula grains.

DISCUSSION

Diurnal cycle — foraging strategies. — In
this study, C. cunicitlarius started its pro-
visioning cycle much earlier than A. vaga.
The specified activity times are probably
dependent on weather conditions. This
earlier departure may have been caused
by higher soil temperatures. However, the
departure time is not known from the oth-
er study site at the Fliegenberg. The in-
vestigated aggregation of C. cunicularius is
exposed southward, has a strong slope
and the soil is only sparsely covered with
vegetation. The exposition of A. vaga's ag-

Table 4. Classification of pollen types other than Salix in the cells of C. cunicularius in 1996.

Family

Genus or species

Percentage [%]

Rosaceae

Fagaceae

Caprifoliceae

Ranunculaceae

Celastraceae

Aceraceae

Sorbus aucuparia, Prunus padus, Primus lau-

5-92

rocerasus, Primus sp., Pyrus sp., Mains sp.

Pilipendula sp.

<1

Quercus sp.

2-10

Sambucus niger

7

Ranunculus sp.

0.2-3.3

Euonymus europaeus

2

Acer sp.

<1

Volume 12, Number 2, 2003

229

Salix □ Acer 0 Prunus □ Quercus

100

-90
n

| 80

S 70

o

o- 60H

§ 50
i 40
a 30

D)

1 20
| 10

I a 5
Pollen load

Fig. 5. Numerical (a) and volumetric (b) percentage of Salix pollen and other pollen types in excavated cells
of C. cwiicularius in the year 1996 and 1998.

gregation is south-eastward, slope is low-
er than in the C. cuniciilarius aggregation
and the sand path is covered with grass.
The surrounding site of A. vaga's aggre-
gation has also more and higher vegeta-
tion cover. Thus, higher soil temperature
at the C. cuniciilarius aggregation may be
caused by these local differences of expo-
sure, slope and vegetation cover (cf. Bis-
choff 2000). Ideally, the diurnal cycle of C.
cuniciilarius should be investigated at ag-
gregations close to A. vaga's aggregation,
i.e. with similar conditions of soil, expo-
sure, slope and vegetation cover.

Several authors have found correlations
between the behavior of bees and weather
conditions (e.g. Linsley 1958, Michener
and Rettenmeyer 1956, Willmer 1983).
Temperature thresholds for the bee's ac-
tivities depend on weather conditions and
the season in which the species occurs.
Many vernal bees begin flight activity at
10 °C and they are less influenced by
cloud cover or wind. Flight activity tem-
peratures reported for other European ear-
ly spring species of Andrena, for instance
A. barbilabris (Kirby), A. cineraria (Linnae-
us), A. clarkclla (Kirby), and the Nearctic
species A. erythronii Robertson, A. vibur-
nella Graenicher, and A. vicina Smith range

from 10 to 16 °C (Gebhardt and Rohr 1987,
Johnson 1981, Michener and Rettenmeyer
1956, Miliczky and Osgood 1995, Stephen
1966, Witt 1992). For vernal species of Col-
letes like C. inaequalis Say and C. validus
Cresson, similar temperature thresholds
are known (Batra 1980). In our study, the
two species began flight activity at 8-9.5
°C air temperature. Similarly, Schonitzer
and Klinksik (1990) recorded flight activi-
ty at a temperature of 8 °C for A. nyc-
theincra Imhoff. Due to unstable weather
conditions in spring, vernal species have
to use days with good weather conditions
very efficiently. To illustrate, on days with
optimal weather conditions provisioning
activity of A. clarkclla is completed in 4 or
5 days (Friese 1923). Late summer species
such as Panurgus banksianus (Kirby) and
Dasypoda hirtipes (Fabricius) often need
temperatures > 20 °C to start their first
trip (Lind 1968, Munster-Swendsen 1968).
Though the flight of A. vaga may already
start at 8 °C, time of its first departure is
later than that of many of the other species
of Andrena mentioned above. This may be
caused by local and seasonal differences in
temperature compared to the other stud-
ies mentioned above. In the present study
the required temperature threshold of 8 °C

230 Journal of Hymenoptera Research

at the investigated aggregation of A. vaga Miliczky and Osgood 1995, Schonitzer

was not reached before 09.00 a.m. This fact and Klinksik 1990).

may explain the strange correlation of the A second reason for the marked differ-
first departure of A. vaga with the temper- ence in daily activity of the two investi-
ature, which represents in fact no corre- gated species could result from A. vaga's'
lation with the temperature. The bees can prolonged stay in the nest. It could not be
start their first trip at a temperature clarified whether A. vaga uses these peri-
threshold of 8 °C and the regression rep- ods in the nest for digging, since no new
resents only the increasing number of sand was pushed to the surface. The oc-
starting bees with time. currence of sand output depends of the se-
Apart from differences in soil tempera- quence of nest construction. When the bee
ture, the beginning of flight activity of the first digs the main burrow and constructs
two species may be influenced by their re- the cells regressively (i.e., the lowest one
spective thermoregulatory abilities. Larger is built first and each subsequent one is at
bees are more likely to achieve flight tern- a higher level), it can fill the inferior main
peratures at low ambient temperatures burrow with the material of the side bur-
(Michener and Rettenmeyer 1956, Stone et rows. Thus no new sand needs to be
al. 1988, Stone and Willmer 1989, Stone pushed out. This has been described by
1993a, b, Stone 1994, Stone et al. 1995, Malyshev (1927) for C. cunicularius and
Wolda and Roubik 1986). Colletes cunicu- also by Rajotte (1979) for C. validus. Yet, A.
larius is one of the largest bees in Germa- vaga constructs its nest conversely, subse-
ny, having a mean heating rate of 7.35 °C quent cells lie deeper and the oldest cell is
per minute (Stone and Willmer 1989). nearest to the surface. Side burrows of A.
These authors investigated the heating- vaga are also filled with sand and in order
rate among A. clarkella and A. fulva (Mull- to fill the first side burrow of a completed
er). These two species are comparable to cell, the bee may use the material of a sec-
A. vaga in body size, hairiness and flight ond side burrow. This was also assumed
season, and differ only in color from A. by Michener and Rettenmeyer (1956) for
vaga. Mean heating rate of these two spe- A. erythronii. Our (Bischoff 2001) and Ma-
cies of Andrena is about 4 to 6.2 °C per lyshev's (1926) description of A. vaga's
minute, respectively. Although C. cunicu- nest architecture are contradictory to the
larius is larger and more hairy than A. descriptions and figures of Friese (1882,
vaga, the abdomen of the latter is deep 1923), in which the last cell is located at
black and passive heat absorbency may be the lowest level. Michener and Rettenmey-
increased. Nevertheless, body size is prob- er (1956) suggested that Friese's nest fig-
ably more important for warming up in ure with cells close along the main burrow
the morning and may enable C. cunicular- like a cluster of grapes resulted from a
ins to begin earlier with daily activity. Af- mixture of different nests lying very close
ter sunset, this species may also benefit together.

from its larger size. In many species, digging of side bur-
Most of the flights of A. vaga took be- rows and new cells has been described to
tween 1 and 2.5 hours on pollen days, and take place in the afternoon (Lind 1968,
between 2 and 3.5 hours on nectar days. Munster-Swendsen 1968, Gebhardt and
The known duration of provisioning trips Rohr 1987, Michener and Rettenmeyer
of other species of Andrena ranged from 20 1956, Witt 1992). However, our results do
minutes to 4 hours, and the number of not confirm these findings for A. vaga,
provisioning trips per day showed a tran- where even after the last provisioning trip,
sition from 1 to 5 (Gebhardt and Rohr females did no intensive digging as ob-
1987, Michener and Rettenmeyer 1956, served in C. cunicularius. Yet, at the begin-

Volume 12, Number 2, 2003

231

Table 5. Results of pollen counts with the electronic counter, the counting chamber and the Lycopodium-
ratio-method (means of all counts of both species).

Grain number/pollen load

Sta

ndard deviation

Mean difference to
electronic counter

Electronic

1053796

722183

counter

1777806

626778

Counting

798069

711542

255728

chamber

Lycopodium-

1850200

515579

498713

ratio-counting

ning of the season, when the aggregation
of A. vaga develops, new tumuli could be
found early in the morning. Thus, we as-
sume that the construction of these tumuli
took place at night or early in the morn-
ing, because sand often was still moisted
and no sand output was observed in the
late evening of the previous day. Andrena
erythronii digs in the late afternoon and
even in the dark (Michener and Retten-
meyer 1956). Nocturnal digging activity
has been reported from other Nearctic
species of Colletes (Batra 1980, Rajotte
1979). Since C. cunicularius constructs its
cells regressively as described above, no
large sand output should occur after the
construction of the main burrow. Digging
activities in the evening can be interpreted
as constructions of new nests, since C. cun-
icularius makes 2 or 3 nests in its life. Col-
letes cunicularius defers the construction of
new nests to the evening time, thus it can
use the whole day for provisioning trips
and can increase the number of construct-
ed nests.

The third explanation for the difference
in daily activity may be the different pol-
len carrying capacities of both species.
Braue (1916) and Friese (1923) described
the different pollen collecting apparatus of
bees and inferred from these differences
the systematic order of bee genera. Since
these early studies, many authors worked
on different pollen collecting apparatus
(Grinfeld 1962, Michener et al. 1978, Pas-
teels and Pasteels 1979, Proctor et al. 1996,
Thorp 1969, Westerkamp 1987). According

to Braue (1916), Andrena is the genus that
can carry home the largest amount of pol-
len with its hind leg brushes and parts of
the thorax. Although both species have
trochanter-femur baskets, the floccus and
thoracic pollen baskets of A. vaga seem to
be more strongly developed than in C.
cunicularius. In the present investigation,
A. vaga collected nearly twice as much
pollen per load on average than C. cuni-
cularius. Exact quantitative data on num-
ber of pollen grains per pollen load are
scarce in the literature. In most cases, only
percentages of different pollen types are
presented, e.g. pollen loads of Andrena
(Chambers 1946). Parker (1981) analyzed
pollen loads of polylectic and oligolectic
bee species quantitatively in order to de-
termine the effectiveness of these species
in pollinating sunflowers. He demonstrat-
ed that female oligoleges carried more
pollen than did any other group of bees
studied. In our study, species collected
much more grains per load than in Park-
er's example; however, the number of car-
ried pollen grains depends highly on the
mean size of the grain type. Accordingly,
our results can only be compared quanti-
tatively to data of the same bee species,
collecting the same pollen species. One
possible factor as to why C. cunicularius
pollen loads were smaller than A. vaga's is
that C. cunicularius collects nectar along
with pollen in each trip. From honey bees
and bumble bees it is known that they can
carry an amount of nectar from 50 to 90%
of their body weight (Heinrich 1979). We

232

Journal of Hymenoptera Research

500-

250-

(0

o
o

t:

(0

o 500

x

Cap 150 Ljm

Range: 13.6 to 80.0 pm
Sample 3 x 400 lI
Counts /ml 5136

25

50

75

pm

250

Cap:

150 urn

Range

13,6 to 80,0 urn

Sample

3x400 pi

Counts / ml

6252

'■ «*-■■*-■-

50
Particle size

75

pm

Fig. 6. Quantitative analysis of the pollen loads of C. cunicularius with the cell counter. Upper graph: 1.
female, 23.04.99, below: 2. female, 27.94.99 (Cap = capillary, particles beneath dotted line = smaller than 10
(xm, no pollen).

were unable to find precise data in the lit- though contains a lot of nectar (Malyshev

erature on the amount of nectar collected 1936). Other bees like Osmia ritfa (Linnae-

by solitary bees on one single trip, but for us) make more dry provisions (Westrich

C. cunicularius it is reported that the pro- 1990).

vision in the cell is extremely liquid and Dividing the grain number of the pure

Volume 12, Number 2, 2003

233

Salix cells for the year 1996 by the grain
number of pure Salix pollen loads (same
year), C. cunicularius had to make approx-
imately nine provisioning trips per cell. In
1998, we had only one cell of C. cunicular-
ius, containing only Salix grains. In this
case, C. cunicularius had to collect seven
pollen loads for completing one cell. If A.
vaga had to gather approximately the
same amount of pollen per cell, it would
have to make only four trips per cells, be-
cause of its bigger carrying capacity per
load. These assumptions agree well with
the observed trips of both species during
one day. Vleugel (1947) observed for A.
vaga only 1 or 2 trips on days with good
weather conditions. The foraging statistics
for Andrena complexa visiting Ranunculus
for pollen show a time of Wi hours to
complete a load, and a pollen foraging
rate of three loads per day (Linsley and
MacSwain 1959). Futhermore the amount
of pollen per cell may depend on the sex
of the offspring. Gerber and Klostermeyer
(1970) provided evidence that females de-
termine the sex of their offspring by fer-
tilizing the egg or not. Males are often
smaller than females and therefore the
stored pollen mass for males is smaller
(Helms 1994, Strickler 1982, Maddocks
and Paulus 1987, Johnson 1988). Regard-
less of the sex of the offspring, it seems
likely that C. cunicularius has to make
more provisioning trips per cell, because
of its smaller pollen carrying capacity. In
conclusion, A. vaga can carry more pollen
per collecting trip; however, due to its
body size C. cunicularius is more indepen-
dent of weather conditions and can be ac-
tive for longer periods per day. Indeed,
the last point is of considerable impor-
tance for vernal bee species, because
weather conditions are often quite unfa-
vorable during spring. Additionally, C.
cunicularius uses the evening and perhaps
the night for digging activity. Our inves-
tigations of daily collecting capacity as
well as those of nest excavations (cf. Bis-
choff 2001) indicate a higher reproduction

rate for C. cunicularius in comparison to A.
vaga.

Nectar- and pollen provisioning trips. — We
observed a rhythm of nectar and pollen
provisioning trips and assumed that A.
vaga collected first all the pollen for one
cell, then added the nectar. Friese (1923)
described exactly this type of behavior for
A. vaga. Dasypoda hirtipes, Andrena ery-
thronii and various species of Anthophora
are also known to add nectar only after
several pollen loads have been carried into
the cell (Lind 1968, Michener and Retten-
meyer 1956, Muller 1884, Westrich 1990).
Malyshev (1936) states that pollen pre-
dominates in the first load or even makes
up the whole load and that the last load
deposited in the cell usually consists
mainly of honey. Other species clearly al-
ternate nectar- and pollen provisioning
trips, e.g. Osinia adunca (Panzer), Osmia
fulviventris Panzer or Chelostoma florisomne
(Linnaeus) (Brechtel 1986, Kapyla 1978,
Westerkamp 1978, Westrich 1990). It is
possible that digger bees and carpenter or
mason bees differ with respect to this be-
havior. Miliczky and Osgood (1995) de-
scribed four trips for A. vicina during
which no pollen was collected (in com-
parison with 64 pollen-collecting trips)
and interpreted them as adult feeding
trips. Since we did not analyze quantity of
nectar in A. vaga's cells, it cannot be defi-
nitely clarified whether nectar trips are
adult feeding trips or nectar provisioning
trips for the offspring. Assuming that A.
vaga cells contain nectar, then there must
be a rhythm between nectar and pollen
provisioning trips. The two observed
flight patterns may represent the different
provisioning behavior for female and
male cells. However, in order to prove this
hypothesis, a longer series of provisioning
trips of a greater number of females have
to be documented and the mass of provi-
sioned pollen has to be analyzed for sex
specifically.

Another reason for the observed pattern
of pollen and nectar trips may be the in-

234 Journal of Hymenoptera Research

fluence of weather conditions. Our under- flowers than Salix, but their reports con-
lying hypothesis was that females make a tain no precise information whether these
nectar trip after a particular hot and dry flower visits were for collecting pollen or
day to increase humidity inside the cell. In nectar. In fact it is not proved at all that
fact, Stephen (1966) noted that the tern- C. cunicularius is really oligolectic on Salix.
perature at which flight activity was ini- Therefore it is not clear whether the col-
lated in A. viburnella was a function of lection of other pollen, as observed in this
weather conditions of the previous day. study, is a result of resource restriction.
Yet, we did not find any correlation of oc- The correlation of increasing percentages
currence of nectar trips to the climatic pa- of other pollen types in the pollen loads
rameters of the previous day. On the con- of C. cunicularius with the end of flowering
trary, after a bad weather day (no flying time of Salix may be an indication for a
conditions the whole day) more females resource limitation. We registered all Salix
made a nectar trip. This may be caused by trees within a radius of 3 km, most of
an mcreased energy consumption after which were S. caprea. In the years 1996 to
one day in the nest. Probably females pro- 1998, these trees were blooming very early
vide themselves with nectar during their and had ceased to flower before females
pollen collecting trips (male Salix plants of C. cunicularius and A. vaga began to col-
also produce nectar). lect pollen. Only several bushes of S. au-
Oligolecty. — Qualitative comparison of ricula were available during nest provi-
collected pollen of both bee species indi- sioning time of both species. To prove
cated important differences in diet whether C. cunicularius collects only other
breadth between the two species: A. vaga pollen when Salix is not available, corn-
collected almost pure Salix pollen, where- parative studies with resource quantifica-
as C. cunicularius collected also a high per- tions at other locations from different
centage of other pollen types. The fact that years are needed.

whole cells contained exclusively other In conclusion, C. cunicularius can not be

pollen types indicates that females of C. regarded as an oligolectic species. The use

cunicularius systematically collect pollen of other, longer blooming host plants,

from other host plants. Early flowering, which are more abundant in the study

tree-like Rosaceae in particular seem to be area, may increase the reproductive suc-

of great importance to this bee species, cess of this species. In contrast to C. cuni-

Mader (1999) listed a number of species of cularius, A. vaga seems not to be affected

Colletes having a relationship to Rosaceae. by the problem of long searching times for

In fact, the Nearctic C. thoracicus Smith pollen sources, since it collected only Salix

and C. nigrifrons Titus are even specialized pollen. However, the activity time of A.

on Rosaceae. In our study, we found also vaga ceased approximately 4 weeks before

pollen loads containing only Quercus, Acer that of C. cunicularius and the problem of

or Ilex pollen, indicating that not only Ro- pollen availability probably did not yet oc-

saceae can replace missing Salix plants. On cur.

the Turkish coast, C. cunicularius females Niche differentiation.— A. vaga and C.
were observed foraging on Pistacia; Salix cunicularius use the same host plant. This
did not occur at this location. In Italy, C. overlap may result in interspecific com-
cunicularius females were observed exclu- petition, if resources are limited. Since
sively on Fabaceae (Kuhlmann in litt). availability of specific pollen is one of the
The whole complex of species, subspecies most relevant niche parameters for bees
and their host-plants seems not yet clear. (Eickwort 1973, Strickler 1979), interspe-
Mader (1999) cited several authors which cific competition in case of a resource re-
observed C. cunicularius on many other striction seems very likely. Niche differ-

Volume 12, Number 2, 2003

235

entiation is often the basis for the coexis-
tence of competitors. There are a number
of ways in which niches can be differen-
tiated. In this case the niches of the two
species seemed to be differentiated on the
basis of conditions. This means that they
use the same resource but their ability to
do so is influenced by environmental con-
ditions and they respond differently to
these conditions (Begon et al. 1990). The
two species show diurnal differences in
their foraging behavior. This temporal
separation is influenced by climatic pa-
rameters such as temperature.

Whether C. cunicularius uses other host
plants because of resource restriction and/
or competition with other species (besides
A. vaga two other Andrena species special-
ized on Salix occur in the study area) or
whether it is not oligolectic at all can only
be proved with removal experiments and
manipulation of the resource availiability.

ACKNOWLEDGMENTS

We thank Annemarie Gossmann, Dr. Beate Kubitz
and Dr. Martina Stebich (Institute for Palaeontology
of the University of Bonn), for introducing us to the
methods of pollen acetolysis and pollen quantifica-
tion with Lycopodium spores. We are grateful to Dirk
Schiffler and Andre Hamm (Institute for Agricultural
Zoology and Bee-Research), for providing a cell coun-
ter and an introduction to this method. We would
like to thank Prof. Dr. G. Vorwohl for determination
of pollen types other than Salix. Special thanks also
to Dr. Rainer Hutterer, Dr. Anrje Bischoff and Dr.
Bradley Sinclair, who reviewed the manuscript. We
would like to thank dozens of students who assisted
during the field work. We are grateful to Prof. Dr. C.
M. Naumann for supporting this study. This work
was funded by a Ph.D. grant from the Deutsche Bun-
desstiftung Umwelt to the senior author.

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J. HYM. RES.
Vol. 12(2), 2003, pp. 238-271

Taxonomic Revision of the Genus Sesioctonus Viereck
(Hymenoptera: Braconidae: Agathidinae)

Rosa A. Briceno G.

Universidad Centroccidental "Lisandro Alvarado", Decanato de Agronomia,

Dpto. de Ciencias Biologicas-Seccion Entomologia, Tarabana, Cabudare,

Estado Lara, Venezuela, e-mail: rabricen@telcel.net.ve

Abstract. — The Neotropical genus Sesioctonus Viereck and its type species S. parathyridis Viereck,
are redescribed. Descriptions for twenty-six new species are presented: S. acrolophus, S. amazo-
nensis, S. ammosakron, S. analogus, S. areolatus, S. ariasi, S. armandoi, S. biospleres, S. boli-
viensis, S. brasiliensis, S. chaconi, S. chrestos, S. clavijoi, S. diazi, S. dichromus, S. dominions, S.
eumenetes, S. galeos, S. garciai, S. grandis, S. kompsos, S. miyayensis, S. peruviensis, S. qui, S.
theskelos, and S. venezuelensis. Seventy-six morphological and color characters are discussed. A
key to species is presented.

Sesioctonus Viereck 1912 is a Neotropical
genus of the subfamily Agathidinae that,
together with the genus Earinus Wesmael,
comprise the tribe Earinini (Sharkey 1992).
Viereck proposed Sesioctonus for the
unique species described so far: Sesiocton-
us parathyridis Viereck.

The main characters that diagnose Se-
sioctonus are: tarsal claws simple, and no-
tauli absent (Viereck 1912; Sharkey 1997).
Many members of Sesioctonus have showy
color patterns with body size varying be-
tween 3.0-12.0 mm. Despite the vivid col-
or patterns and relatively large size, this
genus is poorly represented in insect col-
lections, and there is little biological infor-
mation. Most species of Sesioctonus have
been collected from November until
March and between 100-2000 meters
above sea level. Almost all specimens
were collected in Malaise traps.

All known agathidines are parasitoids
of Lepidoptera larvae (Sharkey 1988,
1997). The only previously known species
of Sesioctonus, S. parathyridis, is recorded
as a larval parasitoid of Parathyris perspi-
cilla Stall (Lepidoptera: Arctiidae), al-
though precise biology data (life cycle) are

not known (Viereck 1912, 1914). Sharkey
(1997) estimated that this genus might in-
clude about thirty species and this revi-
sion includes a total of twenty-seven spe-
cies.

MATERIALS AND METHODS

Specimens are deposited in the follow-
ing collections:

AEI: American Entomological Insti-
tute, Gainesville, FL. USA.

BMNH: The Natural History Museum,
London. England.

CNC: Canadian National Collection,
Biosystematics Research Centre,
Agriculture Canada, Ottawa, On-
tario. Canada.

CUIC: Cornell University Insect Collec-
tion, Department of Entomology,
Cornell University, Ithaca, NY.
USA.

FML: Fundacion Miguel Lillo, Instituto
de Zoologia Miguel Lillo, Tucu-
man. Argentina.

INBio: Instituto de Biodiversidad, Santo
Domingo, Heredia. Costa Rica.

INPA: Instituto Nacional de Pesquisas
da Amazonia. Depto. de Ecologia
e Biologia Evolutiva, Universida-

Volume 12, Number 2, 2003

239

Figs. 1-4. 1, Subpronope triangular in dorso-lateral view. 2, Subpronope oval-shaped in dorso-lateral view.
3-4, Epicnemial carinae. 3, Complete and bilobed. 4, Incomplete and straight.

de Federal de Sao Carlos, Sao
Carlos, SP. Brasil.

MIZA: Museo del Instituto de Zoologia
Agricola "Francisco Fernandez
Yepez", Universidad Central de
Venezuela, Maracay, Aragua. Ve-
nezuela.

MZLU: Museum of Zoology, Lund Uni-
versity, Sweden.

TMB: Termeszettudomanyi Miizeum,
Budapest. Hungary.

UCOB: Museo Entomologico "Jose M.
Osorio", Universidad Centrocci-
dental "Lisandro Alvarado", Tar-
abana, Lara. Venezuela.

UCR: Universidad de Costa Rica, San
Jose. Costa Rica.

UK: Department of Entomology, Uni-
versity of Kentucky, KY. USA.

USNM: United States National Museum,

Smithsonian Institution, Wash-
ington, D. C. USA.
UWY: U. W. Insect Museum, University
of Wyoming, Laramie, WY. USA.
ZSBS: Zoologische Sammlungen des
Bayerischen Staates, Munich, Ger-
many.
The revision was carried out at MIZA at
the Universidad Central de Venezuela in
Maracay, Venezuela.

Several morphometric characters were
used in the descriptions and key. Mea-
surements of Body length (mm) do not in-
clude the ovipositor. Hind femur length and
width (Fig. 5) were measured in lateral
view, with the length taken from the distal
end of the trocantellus to the apex of the
femur. The maximum width was mea-
sured, which is always near the midpoint.
First metasomal tergite length and width re-

240

Journal of Hymenoptera Research

lationship (Fig. 7) refers to the median ter-
gite with the length taken along the mid-
line and the width measured at the apex.
Ovipositor length (mm) was measured from
the base of the hypopygium. In the de-
scription a measurement in parentheses
following a range belongs to the holotype.

The terminology used for the carinae
and areolae of the metanotum and pro-
podeum follows Sharkey (1988), and the
rest of the terminology, including wing
venation follows Sharkey and Wharton et
al. (1997). Additional terms used in this
revision include the occipital tubercles, a
pair of projections, one at each side of the
occipital medial line (Figs. 16-18); and the
median areola of the metanotum, which is the
central area of metanotum often bordered
with carinae (Figs. 25-29).

The key to Sesioctonus species was gen-
erated with the software DELTA version
4.07 (Dallwitz et al. 1997). Seventy six me-
ristic, morphological and color characters
were used for the matrix. The species de-
scriptions also were generated with this
software.

The biology for all the species described
is unknown.

RESULTS AND DISCUSSION

Diversity and distribution of Sesiocton-
us.— The genus is distributed from South-
ern Mexico to Southern Brazil. Members
of Sesioctonus are poorly represented in
entomological collections but most of
these have been collected in Malaise traps
in tropical localities between 0 and 1500
meters above sea level. Only S. areolatus,
has been collected above 2000 meters, spe-
cifically at the Estacion Biologica Las Al-
turas, in Costa Rica.

The poor collecting in many regions of
the Neotropics may explain the scarcity of
this genus in collections. This is clearly
shown by the numerous specimens from
Costa Rica and Panama where collecting
has been extensive in recent years.

Color patterns. — Color patterns (yellow
and black) shown by many braconids, in-

cluding Sesioctonus, are frequently shared
with other orders of insects, specially Co-
leoptera and Hemiptera. However, some
Lepidoptera, Neuroptera and Diptera also
exhibit (Quicke 1997) these color patterns.
Certainly, specimens of Sesioctonus could
be confused with specimens of some of
these groups when they are observed in
nature, but the existence of a large mi-
metic complex has not yet been conclu-
sively demostrated. At taxonomic level,
the various combination of yellow, or-
ange, and black colors on the body of
wasps are important characters for the
separation of species.

Wing color pattern. — The color pattern of
the wings seems to be related to geo-
graphical distribution. Most species from
South America have the wings banded
from the base: yellow, infuscate, yellow,
infuscate; while most species from Central
America have the wings entirely infuscate.
Another curious fact is that the only spe-
cies from an island (S. dominions) has hy-
aline wings. These observations could
suggest that there are mimetic relation-
ships with other insects on the continent,
or perhaps the presence of a sham apose-
matic behavior, which was not present in
the island.

Genus Sesioctonus Viereck

Sesioctonus Viereck 1912:1. Type species: Sesioc-
tonus parathyridis Viereck. (Monobasic and
original designation). Viereck 1914: 133.

Diagnosis. — Sesioctonus species may be
distinguished from other agathidines us-
ing the following combination of charac-
ters: Body smooth and bright, lacking
sculpture, scutellar depression smooth,
notauli absent, tarsal claws simple (Fig.
32), epicnemial carina bilobed between
fore coxae; rarely (8%) straight, ovipositor
about as long as body length.

Description. — Head: Antenna with 23-48
flagellomeres, usually with 28-35; inter-
antennal space with distinct keel (14%), or
flat without distinct keel; antennal sockets

Volume 12, Number 2, 2003

241

^

Foretibial
spines

7^

first metasomal
median tergite width

Figs. 5-8. 5, Hing legs of Braconidae (Wharton et al. 1997) showing measurements done. 6, Apex of foretibia
with spines. 7, First metasomal median tergite showing measurements done. 8, Propodeum, a, with central
areola; b, without areola.

242

Journal of Hymenoptera Research

Figs. 9-10. Venation in Sesioctonus. 9, Forewing, a, (RS+M)a vein incomplete; b, 3RSa vein, hind wing; c,
Cub vein not tubular. 10, Forewing, a, (RS+M)a vein complete.

excavated (51%); face rarely with longitu-
dinal median carina; mandibles with two
teeth, usually the outer tooth as long as
the inner; maxillary palpus with four or
five segments; labial palpus with three or
four segments; third segment of labial pal-
pus, when present, shorter and partially
fused to fourth segment; gena usually
moderately expanded posteroventrally
(37%), sometimes not expanded; occipital
tubercles often present (51%); occiput usu-
ally excavated. Mesosoma: Smooth and
bright, without sculpture; subpronope tri-

angle-shape or oval-shape; notauli always
absent; scutellum generally convex in lat-
eral view, rarely flat; lateral carina of the
scutellar depression usually absent; me-
dian areola of the metanotum often with
longitudinal carina and lateral carinae that
join posteriorly or not, generally smooth,
sometimes with small rugosities; propo-
deum convex (49%) or flat (51%) in lateral
view; longitudinal carina of propodeum
usually absent, sometimes present; epic-
nemial carina absent (6%) or present
(94%), complete (70%) or incomplete lat-

Volume 12, Number 2, 2003

243

erally (24%), generally bilobed medially
between fore coxae, sometimes straight;
hind coxal cavities open, forming a com-
mon foramen with the metasoma; fore
wing (RS+M)a vein present, complete
(57%) or incomplete (43%); fore wing 3RSa
usually present; hind wing with 3-10
hamuli, generally 4-6; hind wing 2-1A
vein usually tubular; hind wing CUb vein
not tubular; fore tibia sometimes with

spines; medial tibia usually with 2-16
spines; hind tibia with 5-25 spines, usu-
ally 12-17; tarsal claws simple on all legs.
Metasoma: Smooth and bright, without
sculpture; median tergite of the first me-
tasomal segment usually with two longi-
tudinal dorsal carinae, rarely with pits
posterad spiracle; ovipositor as long as
length of the body. Length. 2.0-12. 0mm;
excluding ovipositor.

KEY TO SPECIES OF SESIOCTONUS

1 Occipital tubercles present (Figs. 16-18) 2

Occipital tubercles absent (Figs. 19) 13

2(1) Epicnemial carina straight medially or absent (indented at midline, between forecox-

ae), sometimes difficult to see (Figs. 4, 23) 3

Epicnemial carina bilobed medially, (indented at midline, between the forecoxae) (Figs.
3, 22) 5

3(2) Epicnemial carina complete in lateral view (Figs. 3, 22) garciai sp. n.

Epicnemial carina incomplete or absent in lateral view (Fig. 23) 4

4(3) Face with median longitudinal carina (Fig. 13) acrolopilms sp. n.

Face without median longitudinal carina (similar to Figs. 12, 14) analogns sp. n.

5(2) Midcoxa not completely melanic, color variable 6

Midcoxa completely melanic 7

6(5) Fore tibia with spines (Figs. 6, 30), midcoxa yellowish orange, forewing (RS+M)a vein

complete (Fig. 10a) and 3RSa vein present (Fig. 9b) peruviensis sp. n.

Fore tibia without spines, midcoxa melanic dorsally and yellowish orange ventrally,

forewing (RS+M)a vein complete (Fig. 10a) and 3RSa vein absent (Fig. 10)

chaconi sp. n.

7(5) Longitudinal carina of scutellar depression present venezuelensis sp. n.

Longitudinal carina of scutellar depression absent 8

8(7) Median areola of metanotum with longitudinal rugosities (Fig. 29), median tergite of
first metasomal segment without pair of lateral longitudinal carinae (similar to Fig.

34), fore wing (RS+M)a vein complete (Fig. 10a) kompsos sp. n.

Median areola of metanotum smooth (Figs. 25-28) 9

9(8) Mesoscutum melanic 10

Mesoscutum yellowish orange 11

10(9) Fore wing infuscate with large hyaline spot, metasoma reddish brown

brasiliensis sp. n.

Fore wing either infuscate without hyaline spot or hyaline basally, infuscate apically,
mesosoma melanic and metasoma yellowish orange dichromus sp. n.

11(9) Median longitudinal carina of propodeum present and complete ariasi sp. n.

Median longitudinal carina of propodeum absent or incomplete 12

12(11) Subpronope triangular (Fig. 1), fore wing 3RSa vein absent (Fig. 10) . . boliviensis sp. n.
Subpronope oval-shape (Fig. 2), fore wing 3RSa vein present (Fig. 9) diazi sp. n.

244 Journal of Hymenoptera Research

13(1) Occiput excavated (similar to Figs. 16-18) eumenetes sp. n.

Occiput not excavated (Fig. 19) 14

14(13) Median areola of metanotum with lateral carinae (Figs. 25-28), flagellum with less
than 40 flagellomeres, interantennal space with a rounded longitudinal keel or keel

absent (Fig. 12), never sharp; specimens with less than 8mm in body length 15

Median areola of metanotum without sharp lateral carinae (Fig. 29), flagellum with 40
flagellomeres or more, interantennal space with a sharp longitudinal keel (Fig. 11),
specimens greater than 8mm in body length 25

15(14) Median areola of metanotum with lateral carinae present and meeting posteriorly

(Figs. 25, 26) 16

Median areola of metanotum with lateral carinae present and not meeting posteriorly
(Figs. 27, 28) 23

16(15) Epicnemial carina present (Figs. 3, 4) 17

Epicnemial carina completely absent ammosakron sp. n.

17(16) Epicnemial carina complete laterally (Fig. 3) 18

Epicnemial carina incomplete laterally (Fig. 4) 21

18(17) Hind tibia melanic amazonensis sp. n.

Hind tibia mostly yellowish orange 19

19(18) Propodeum with central areola absent 20

Propodeum with central areola present (Fig. 8a) areolatus sp. n.

20(19) Flagellum with 32 flagellomeres, interantennal space with rounded longitudinal keel
(similar to Fig. 12), hind tibia yellowish orange in basal half, melanic apically . . .

miyayensis sp. n.

Flagellum with 25 flagellomeres, interantennal space without longitudinal keel, hind
tibia mostly yellowish orange, melanic apically clavijoi sp. n.

21(17) Epicnemial carina straight medially (between forecoxae) (Fig. 4), body length less than

3mm dominions sp. n.

Epicnemial carina bilobed medially (indented at midline, between forecoxae) (Fig. 3),
body length more than 3mm 22

22(21) Forewing (RS + M)a vein complete (Fig. 10a) armandoi sp. n.

Forewing (RS + M)a vein incomplete (Fig. 9a) biospleres sp. n.

23(15) Epicnemial carina present, complete or incomplete laterally (Figs. 3, 4) 24

Epicnemial carina completely absent chrestos sp. n.

24(23) Epicnemial carina straight medially (indented at midline, between forecoxae) (Fig. 4)

galeos sp. n.

Epicnemial carina bilobed medially (indented at midline, between forecoxae) (Fig. 3)

theskelos sp. n.

25(14) Third and fourth labial palpomeres not fused, first metasomal median tergite with

depression posterad spiracle (Figs. 36, 37) grandis sp. n.

Third and fourth labial palpomeres fused, first metasomal median tergite with or with-
out depression posterad spiracle 26

26(25) First metasomal median tergite with depression posterad spiracle (similar to Figs. 36,

37) qui sp. n.

First metasomal median tergite without depression posterad spiracle

parathyridis Viereck

Volume 12, Number 2, 2003

245

Sesioctomis acrolophns Briceno, sp. n.

Figs. 11, 13, 15, 21, 23, 24, 25, 30, 32, 35

Diagnosis. — Face with median longitu-
dinal carina, interantennal space with a
sharp longitudinal keel and median areola
of metanotum with median longitudinal
carina. S. analogous is similar but can be
distinguished by this combination of char-
acters.

Description. — 9. Length. Body, 9-10 mm
(9.5). Head: Flagellum with 32 flagello-
meres. Interantennal space with sharp lon-
gitudinal keel. Antennal sockets deeply
excavated. Face with median longitudinal
carina. Genae strongly expanded poster-
oventrally. Occipital tubercles present. Oc-
ciput excavated. Mandible concave, outer
tooth longer than inner tooth. Maxilla
with 5 palpomeres. Third and fourth labial
palpomeres not fused. Mesosoma: Sub-
pronope oval. Longitudinal carinae of scu-
tellar depression absent. Scutellum con-
vex. Median areola of metanotum smooth;
with median longitudinal carina; and with
lateral carinae present and meeting pos-
teriorly. Propodeum convex, median lon-
gitudinal carina present. Epicnemial cari-
na blunt, incomplete dorsally, straight me-
dially (between forecoxae). Foretibial
spines present. Midtibia with 8 spines.
Hind tibia with 11 spines. Hind femur 3.17
times as long as wide. (RS + M)a vein of
forewing incomplete. 3RSa vein of fore-
wing present. 2-1A vein of hind wing not
tubular. CUb vein of hind wing not tu-
bular. Hind wing with 4-5 hamuli (4). Me-
tasotna: Median tergite of first metasomal
segment with pair of lateral longitudinal
carinae. First metasomal median tergite
without depression posterad spiracle.
Length width ratio of first metasomal me-
dian tergite 1.09. Ovipositor 9-10 mm
(9.2). Color: Head melanic except maxil-
lary and labial palpomeres sometimes yel-
lowish orange. Antenna melanic. Meso-
soma yellowish orange. Forelegs melanic
except coxae yellowish orange. Midlegs
melanic except coxae sometimes yellowish

orange. Hindleg melanic except coxa and
femur mostly yellowish orange, but me-
lanic distally. Forewing yellow basally
and infuscate apically. Stigma melanic, or
yellowish orange. Hind wing yellow ba-
sally infuscate apically. Metasoma yellow-
ish orange. Ovipositor yellowish orange.

6 . — Unknown.

Material examined. — Holotype: Costa
Rica: 9, CRI002 492066, Prov. Alajuela,
sector Colonia Palmarena, 700m, oct.1996,
G. Carballo (INBio). Paratopes: Costa Rica:
9, Heredia, Est. Biol. La Selva, 50-1 50m,
10° 26'N 84° 01 'W, 02 de Marzo 1993,
bosque primario (INBio); 9, R. San Lor-
encito, 900m, R. F. San Ramon, 5km N de
Colonia Palmarena, Prov. Alajuela, 13-18
Junio.1993, (without abdomen) (INBio); 9,
Heredia, 3km S. Puerto Viejo, OTS, La Sel-
va, 100m, xii.1992, P Hanson, (UWY); 9,
Heredia, 3km S. Puerto Viejo OTS, La Sel-
va, 100m. l-15.ix.1992, P. Hanson, huer-
tos, set de trampas malaise de G. Wright
(UCR); 9, La Selva, 15.xii.1993, J Longino
(M/04/272) (UK); 29, Heredia, Est. Biol.
La Selva. 50-150m, 10° 26'N 84° 01' W,
x.1992, P Hanson, C Godoy (UCR)
(UCOB); 9, Limon. 16km W. Guapiles,
400m, i-iv. 1992, col. Paul Hanson (UCR).

Distribution. — Known only from low-
land Atlantic rain forests in Costa Rica, up
to 900m.

Etymology. — From Greek acrolophus that
means keel, in reference to the longitudi-
nal carina on the face in this species.

Sesioctomis amazonensis Briceno, sp. n.

Diagnosis. — S. amazonensis share charac-
ters with S. armandoi from which it is sep-
arated by the lack of longitudinal rugosi-
ties on the median areola of metanotum
and the presence of a complete epicnemial
carina.

Description. — 9. Length. Body 5-7 mm
(7.0). Head: Flagellum with 35-38 flagel-
lomeres (38). Interantennal space with
rounded longitudinal keel. Antennal sock-
ets moderately excavated. Face without
median longitudinal carina. Genae not ex-

246

Journal of Hymenoptera Research

Figs. 11-15. Head. 11, S. acrolophus, arrow shows interantennal space with a sharp longitudinal keel. 12, S.
dichromus in dorsal view showing interantennal space with a rounded longitudinal keel or absent. 13, S.
acrolophus, arrow shows face with median longitudinal carina. 14, S. brasiliensis. 15, S. acrolophus, arrow shows
expanded gena.

panded posteroventrally. Occipital tuber-
cles absent. Occiput not excavated. Man-
dible flat, outer tooth of mandible not
longer than inner tooth. Maxilla with 4
palpomeres. Third and fourth labial pal-

pomeres completely fused. Mesosoma:
Subpronope triangular. Longitudinal ca-
rinae of scutellar depression absent. Scu-
tellum convex. Median areola of metano-
tum smooth; without median longitudinal

Volume 12, Number 2, 2003

247

carina; and with lateral carinae present
and meeting posteriorly. Propodeum con-
vex, median longitudinal carina absent.
Epicnemial carina, sharp, complete, bi-
lobed medially (between forecoxae). Fore-
tibial spines absent. Midtibia with 5-9
spines (9). Hind tibia with 10 spines. Hind
femur 3.51 times as long as wide.
(RS + M)a vein of forewing complete. 3RSa
vein of forewing absent. 2-1 A vein of hind
wing tubular. CUb vein of hind wing not
tubular. Hind wing with 4-5 hamuli (5).
Metasoma: Median tergite of first meta-
somal segment without pair of lateral lon-
gitudinal carinae. First metasomal median
tergite without depression posterad spi-
racle. Length width ratio of first metaso-
mal median tergite 1.01. Ovipositor 7 mm.
Color: Head melanic. Antenna melanic.
Maxillary palpomeres yellowish orange
except basal two palpomeres melanic. La-
bial palpomeres yellowish orange. Meso-
soma yellowish orange. Forelegs yellow-
ish orange except femur sometimes me-
lanic ventrally. Midlegs yellowish orange
except tibia and tarsus melanic. Hindleg
melanic with coxa yellowish orange. Fore-
wing banded from base, yellow, infuscate,
yellow, infuscate. Stigma melanic. Hind
wing banded from base, yellow, infuscate,
yellow, infuscate. Metasoma yellowish or-
ange. Ovipositor yellowish orange.

6 . — Unknown.

Material examined. — Holotype. 9 . Brasil:
Amazonas, Res. Ducke, 26km NE Manaus,
22.07.1981, J.A. Rafael, trampa malaise
(CNC). Paratypcs. BRASIL: 9, same data
as holotype (CNC); 9, Manaus, ZF3, Km
23, Faz.Esteio, Res 1112; B. Klein col.
21.1.1986 (INPA).

Distribution. — This species is known
only from the Amazonas region of Brasil.

Etymology. — This species is named after
the locality of the holotype specimen.

Sesioctonus ammosakron Briceno, sp. n.

Diagnosis. — Epicnemial carina absent, a
characteristic shared with S. chrestos and
S. grandis. However, fore tibial with spines

are present the longitudinal carina of the
median metasomal tergite are absent in S.
chrestos. S. grandis can be separated by its
larger size and the presence of 4 maxillary
palpomeres.

Description. — 9. Length. Body 3.5 mm.
Head: Flagellum with 24 flagellomeres. In-
terantennal space without longitudinal
keel. Antennal sockets not excavated. Face
without median longitudinal carina. Ge-
nae not expanded posteroventrally. Occip-
ital tubercles absent. Occiput not excavat-
ed. Mandible concave, outer tooth of man-
dible not longer than inner tooth. Maxilla
with 5 palpomeres. Third and fourth labial
palpomeres not fused. Mesosoma: Sub-
pronope oval. Longitudinal carinae of scu-
tellar depression absent. Scutellum con-
vex. Median areola of metanotum smooth;
without median longitudinal carina; and
with lateral carinae present and meeting
posteriorly. Propodeum convex, median
longitudinal carina absent. Epicnemial ca-
rina completely absent. Foretibial spines
absent. Midtibia with 9 spines. Hind tibia
with 15 spines. Hind femur 3.14 times as
long as wide. (RS + M)a vein of forewing
incomplete. 3RSa vein of forewing absent.
2-1A vein of hind wing tubular. CUb vein
of hind wing not tubular. Hind wing with
3 hamuli. Metasoma: Median tergite of
first metasomal segment with pair of lat-
eral longitudinal carinae. First metasomal
median tergite without depression poster-
ad spiracle. Length width ratio of first me-
tasomal median tergite 1.32. Ovipositor
3.5 mm. Color: Head yellowish orange ex-
cept vertex and occiput melanic. Antenna
melanic. Maxillary and labial palpomeres
yellowish orange. Mesosoma yellowish
orange sometimes melanic. Forelegs yel-
lowish orange. Midlegs yellowish orange.
Hindleg yellowish orange except tibia and
tarsus melanic distally. Forewing entirely
infuscate. Stigma melanic. Hind wing en-
tirely infuscate. Metasoma yellowish or-
ange but third tergum with median tergite
melanic in posterior quarter; fourth ter-
gum with median tergum melanic and

248

Journal of Hymenoptera Research

fifth to eighth metasomal terga mostly yel-
lowish orange but median tergites melanic
centrally. Ovipositor yellowish orange.

8 . — Unknown.

Material examined. — Holotype. Costa
Rica: 9, Puntarenas, San Vito, Est. Biol.
Las Alturas, 1500m. iii.1992, Paul Hanson.
(UWY).

Distribution. — This species is known
only from the Puntarenas region of Costa
Rica.

Etymology. — From Greek aminos that
means sand and akron that means top, in
reference to holotype locality, Puntarenas.

Sesioctomts analogus Briceno, sp. n.

Diagnosis. — S. analogus can be distin-
guished from most Sesioctonus species by
the presence of 5 maxillary palpomeres
and 4 labial palpomeres. However, this
character occurs in 5 other species of the
genus, of which S. acrolophus is closest to
S. analogus. These two species are separat-
ed by the presence of a longitudinal carina
on the face of S. acrolophus, which is absent
in S. analogus.

Description. — 9. Length. Body 10 mm.
Head: Flagellum with broken after flagel-
lomere 20. Interantennal space with
rounded longitudinal keel. Antennal sock-
ets not excavated. Face without median
longitudinal carina. Genae strongly ex-
panded posteroventrally. Occipital tuber-
cles present. Occiput excavated. Maxilla
with 5 palpomeres. Third and fourth labial
palpomeres not fused. Mesosoma: Sub-
pronope oval. Longitudinal carinae of scu-
tellar depression absent. Scutellum con-
vex. Median areola of metanotum smooth;
without median longitudinal carina; and
with lateral carinae present and meeting
posteriorly. Propodeum convex, median
longitudinal carina present. Epicnemial
carina blunt, incomplete dorsally, straight
medially (between forecoxae). Foretibial
spines present. Midtibia with 18 spines.
Hind tibia with 21 spines. Hind femur 2.88
times as long as wide. (RS + M)a vein of
forewing incomplete. 3RSa vein of fore-

wing present. 2-1 A vein of hind wing tu-
bular. CUb vein of hind wing not tubular.
Hind wing with 4 hamuli. Metasoma: Me-
dian tergite of first metasomal segment
with pair of lateral longitudinal carinae.
First metasomal median tergite without
depression posterad spiracle. Length
width ratio of first metasomal median ter-
gite 0.99. Ovipositor 8 mm. Color: Head
melanic. Antenna melanic. Maxillary and
labial palpomeres yellowish orange. Me-
sosoma yellowish orange. Forelegs melan-
ic sometimes coxa and tarsus yellowish
orange. Midlegs melanic sometimes coxa,
femur and basitarsus yellowish orange.
Hindleg melanic, sometimes coxa, femur
and basitarsus yellowish orange basally.
Forewing yellow basally and infuscate
apically. Stigma melanic. Hind wing yel-
low basally infuscate apically. Metasoma
yellowish orange. Ovipositor yellowish
orange.

6 . — Unknown.

Material examined. — Holotype: Costa
Rica: 9, Est. Pitilla, 9km S, Sta. Cecilia,
A.C., Guanacaste, Prov. Guana, 700m,
v.1994, P. Rios, Malaise (INBio).

Distribution. — This species is known
only from lowlands of Costa Rica up to
700m.

Etymology. — This species name refers to
a similarity with S. acrolophus.

Sesioctonus areolatus Briceno, sp. n.

Diagnosis. — Presence of central areola
on the propodeum, median longitudinal
carina of the scutellar depression, foretib-
ial with spines and epicnemial carina com-
plete and straight medially. Also, the body
size is small.

Description. — 9. Length. Body, 4-5 mm
(4.2). Head: Flagellum with 26-28 flagel-
lomeres (26). Interantennal space without
longitudinal keel. Antennal sockets not ex-
cavated. Face without median longitudi-
nal carina. Genae not expanded poster-
oventrally. Occipital tubercles absent. Oc-
ciput not excavated. Mandible concave,
outer tooth of mandible not longer than

Volume 12, Number 2, 2003

249

Rk>

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Par

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Figs. 16-21. Head. 16-19, Occipital tubercles. 16, 18, S. dichromus. 17, S. diazi. 19, S. grandis without tubercles.
20-21, Mandible teeth. 20, S. dichromus with outer tooth not longer than inner tooth. 21, S. acrolophus with
outer tooth longer than inner tooth.

inner tooth. Maxilla with 4 palpomeres.
Third and fourth labial palpomeres com-
pletely fused. Mesosoma: Subpronope tri-
angular. Longitudinal carinae of scutellar
depression present. Scutellum convex.
Median areola of metanotum smooth;
without median longitudinal carina; and

with lateral carinae present and meeting
posteriorly. Propodeum flat, median lon-
gitudinal carina present. Epicnemial cari-
na sharp, complete, straight medially (be-
tween forecoxae), sometimes bilobed.
Foretibial spines present. Midtibia with 7-
8 spines (7). Hind tibia with 10-12 spines

250 Journal of Hymenoptera Research

(10). Hind femur 3.35 times as long as acters: presence of occipital tubercles, me-
wide. (RS+M)a vein of forewing incom- dian areola of metanotum with lateral ca-
plete. 3RSa vein of forewing present. 2-1A rinae not meeting posteriorly, and
vein of hind wing tubular. CUb vein of presence of the longitudinal carinae in the
hind wing not tubular. Hind wing with 3 propodeum. This combination could con-
hamuli. Metasoma: Median tergite of first fuse S. ariasi with S. acrolophus, however
metasomal segment with pair of lateral these species can be separate easily be-
longitudinal carinae. First metasomal me- cause the number of maxillary and labial
dian tergite without depression posterad palpomere is fewer in S. ariasi.
spiracle. Length width ratio of first meta- Description. — 9. Length. Body, 6-8.5 mm
somal median tergite 1.04. Ovipositor 3- (8.0). Head: Flagellum with 30-34 flagel-
4.3 mm (4.0). Color: Head yellowish or- lomeres (33). Interantennal space with
ange. Antenna melanic. Maxillary and la- rounded longitudinal keel. Antennal sock-
bial palpomeres yellowish orange, some- ets not excavated. Face without median
times melanic. Mesosoma entirely longitudinal carina. Genae not expanded
yellowish orange. Forelegs yellowish or- posteroventrally. Occipital tubercles pre-
ange. Midlegs yellowish orange some- sent. Occiput excavated. Mandible con-
times tibia melanic. Hindleg yellowish or- cave, outer tooth of mandible not longer
ange with tibia yellowish orange but me- than inner tooth. Maxilla with 4 palpo-
lanic distally; and tarsus melanic, or yel- meres. Third and fourth labial palpomeres
lowish orange. Forewing entirely infuscate completely fused. Mesosoma: Subpronope
or infuscate with hyaline spots. Stigma oval. Longitudinal carina of scutellar de-
melanic. Hind wing entirely infuscate. pression absent. Scutellum convex. Medi-
Metasoma yellowish orange. Ovipositor an areola of metanotum smooth; without
yellowish orange. median longitudinal carina; and with lat-

6 . — Unknown. eral carinae present and not meeting pos-

Material examined. — Holotype. Costa teriorly. Propodeum convex, median Ion-
Rica: 9, Puntarenas, San Vito, Est. Biol, gitudinal carina present. Epicnemial cari-
Las Alturas, 2100m, iii-v.1995, Paul Han- na sharp, complete, bilobed medially (be-
son (UWY). Paratypes. Costa Rica: 39, tween forecoxae). Foretibial spines absent.
Puntarenas, San Vito, Est. Biol. Las Altu- Midtibia with 8 spines. Hind tibia with
ras, 1500-2100m, v. 1992, iii-v.1995, Paul 14-16 spines (15). Hind femur 3.32 times
Hanson (UWY); 9, Guanacaste Prov., as long as wide. (RS+M)a vein of fore-
300m, decidious woods (25-30 yrs), 4.ix- wing incomplete. 3RSa vein of forewing
5.x. 1985, Gauld & Janzen (UK). Honduras: present. 2-1A vein of hind wing tubular.
9 , Cortes, Parque Nacional Cusuco 5km CUb vein of hind wing not tubular. Hind
N de Buenos Aires, 15°29'N 88°13'W, wing with 5 hamuli. Metasoma: Median
15.X.1995, malaise trap, R. Cave (MZLU). tergite of first metasomal segment without

Distribution. — This species is known pair of lateral longitudinal carinae. First

only from Costa Rica and Honduras. metasomal median tergite without depres-

Etymology. — From Latin areolatus that sion posterad spiracle. Length width ratio

means areolated, in reference to the cen- of first metasomal median tergite 1.18.

tral areola on the propodeum in this spe- Ovipositor 5-8 mm (7.5). Color: Head me-

cies- lanic. Antenna melanic. Maxillary pal-

c • x ■ n • ~ pomeres melanic, sometimes yellowish or-

beswctonus ariasi Bnceno, sp. n. . , , J ,

r ange except basal two palpomeres melan-

Diagnosis.—S. ariasi can be distin- ic. Labial palpomeres melanic. Mesosoma

guished from the rest of Sesioctonus spe- mostly yellowish orange with pronotum

cies by the following combination of char- melanic anteriorly, propleuron and meta-

Volume 12, Number 2, 2003 251

pleuron melanic, or yellowish orange, latter species is straight and not bilobed as

sometimes melanic posteriorly. Forelegs S. armandoi.

melanic except tibia melanic, or yellowish Description. — 9. Length. Body, 6 mm.

orange and tarsus yellowish orange. Mid- Head: Flagellum with 34 flagellomeres. In-

legs melanic except tarsus with basitarsus terantennal space with rounded longitu-

sometimes yellowish orange in basal half, dinal keel. Antennal sockets moderately

Hindleg melanic. Forewing entirely infus- excavated. Face without median longitu-

cate. Stigma melanic. Hind wing entirely dinal carina. Genae not expanded poster-

infuscate. Metasoma mostly yellowish or- oventrally. Occipital tubercles absent. Oc-

ange except fifth to eighth metasomal ter- ciput not excavated. Mandible concave,

ga melanic. Ovipositor yellowish orange Maxilla with 4 palpomeres. Third and

except apical eighth melanic. fourth labial palpomeres completely

6\ — Essentially as female. fused. Mesosoma: Subpronope triangular.

Material examined. — Holotype: Brazil: 9, Longitudinal carinae of scutellar depres-

Matogrosso, 12° 31 S, 55° 37 W, ii. 1976, sion absent. Scutellum convex. Median

M. Alvarenga (CNC). Parah/pes: Colom- areola of metanotum with longitudinal ru-

bia: 8, Cundinamarca, Monterredondo, gosities; without median longitudinal ca-

14.xii.1958, J. Foerster (CNC), 9, Antio- rina; and with lateral carinae present and

quia, Mun. San Luis R.N. Rio Claro, El Re- meeting posteriorly. Propodeum convex,

fugio 5° 47'N, 75° O'W, 500m, malaise, median longitudinal carina absent. Epic-

13.i.98. Diego Campos (UK); 9, Amazon- nemial carina, sharp, incomplete dorsally,

as, PNN. Amacayacu, Mocagua, 3° 23'S, bilobed medially (between forecoxae).

70° 06'W, 150m, malaise, 26.ii-12.iii.2001, Foretibial spines absent. Midtibia with 5

B. Amado (UK); 9, Cauca, PNN. Gorgona, spines. Hind tibia with 10 spines. Hind fe-

Mancora, 2° 58' N, 78° 11' W, 60m, mal- mur 3.27 times as long as wide. (RS+M)a

aise, 26.vi-18.vii.2000, H. Torres (UK), vein of forewing complete. 3RSa vein of

Costa Rica: 8 , Est. Hitoy Cerere, 100m, R. forewing absent. 2-1 A vein of hind wing

Cerere, Res. Biol. Hitoy Cerere, Prov. Li- tubular. CUb vein of hind wing not tu-

mon, vii.1992, G. Carballo (INBio). Bolivia: bular. Hind wing with 4 hamuli. Metaso-

9 , Staudinger K (ZSBS); Ecuador: 8 , ma: Median tergite of first metasomal seg-

Napo, Prov. Sacha, 9.iii.l983, L. Huggert ment without pair of lateral longitudinal

(CNC). carinae. First metasomal median tergite

Distribution. — From Costa Rica in Cen- without depression posterad spiracle,

tral America to Brazi, Bolivia and Colom- Length width ratio of first metasomal me-

bia in South America. dian tergite 0.94. Ovipositor 6 mm. Color:

Etymology. — This species is named in Head melanic. Antenna melanic. Maxil-

honor of Quintin Arias, for his friendship lary and labial palpomeres yellowish or-

and advise regarding computer software. ange. Mesosoma melanic except metano-
tum, propodeum and metapleuron yel-

Sesioctonus armandoi Briceno, sp. n. }owish omnge Forelegs yeliowish orange.

Diagnosis. — S. armandoi can be distin- Midlegs yellowish orange. Hindleg me-

guished from the rest of Sesioctonus spe- lanic except trochanter and trocantellus

cies by the following combination of char- yellowish orange. Forewing banded from

acters: occipital tubercles absent, epicne- base, yellow, infuscate, yellow, infuscate.

mial carina incomplete laterally, median Stigma melanic. Hind wing banded from

areola of metanotum with longitudinal ru- base, yellow, infuscate, yellow, infuscate.

gosities and with lateral carinae meeting Metasoma yellowish orange until third

posteriorly. This combination is present in tergum, rest melanic. Ovipositor yellowish

S. galeos but the epicnemial carina in this orange.

252

Journal of Hymenoptera Research

Figs. 22-27. 22-23, Epicnemial carina. 22, Complete and bilobed in S. brasiliensis, 23, incomplete and straight
in S. acrolophus. 24, S. acrolophus in lateral view, right arrow shows longitudinal carina of propodeum and left
arrow first metasomal segment with pair of lateral longitudinal carinae. 25-27, Median areola of metanotum.
25, Smooth with longitudinal carinae and lateral carinae meeting posteriorly in S. acrolophus. 26, Smooth with
lateral carinae meeting posteriorly in S. clavijoi. 27, Smooth with lateral carinae not meeting posteriorly in S.
brasiliensis.

6\— Essentially as female. N, 70° 06' W, 150m, 7-19.vii.2000, A Par-

Material examined. — Holotype: Ecuador: ente (UK); S , Amazonas, PNN Amacaya-

9, Napo & Coca Rivers, 2-10.V.1965, Luis cu Matamata, 8-12.iii.2000, Sharkey (UK).

Pena (AEI). Paratypes: Colombia: $,Ama- Distribution. — This species is known

zonas, PNN Amacayacu Mocagua, 3° 23' only from Ecuador and Colombia.

Volume 12, Number 2, 2003

253

Etymology. — This species is named in
honor of Armando Briceno, Venezuelan
entomologist, and my uncle.

Sesioctonus biospleres Briceno, sp. n.

Diagnosis. — S. biospleres can be distin-
guished by the following combination of
characters: genae expanded posteriorly,
occipital tubercles absent, epicnemial ca-
rina incomplete laterally, foretibia with
spines and median tergite of first metaso-
mal tergum with pair of longitudinal ca-
rinae.

Description. — 9. Length. Body, 6-8 mm
(6.5). Head: Flagellum with 30-35 flagel-
lomeres (34). Interantennal space with
rounded longitudinal keel, or keel lacking.
Antennal sockets not excavated. Face
without median longitudinal carina. Ge-
nae expanded posteroventrally. Occipital
tubercles absent. Occiput not excavated.
Mandible concave, outer tooth of mandi-
ble not longer than inner tooth. Maxilla
with 4 palpomeres. Third and fourth labial
palpomeres completely fused. Mesosoma:
Subpronope triangular. Longitudinal ca-
rinae of scutellar depression absent. Scu-
tellum convex. Median areola of metano-
tum smooth; without median longitudinal
carina; and with lateral carinae present
and meeting posteriorly. Propodeum con-
vex, median longitudinal carina present.
Epicnemial carina, sharp, incomplete lat-
erally, bilobed medially (between forecox-
ae). Foretibial spines present. Midtibia
with 6-10 spines (9). Hind tibia with 10-
14 spines (14). Hind femur 3.11 times as
long as wide. (RS + M)a vein of forewing
incomplete. 3RSa vein of forewing absent.
2-1 A vein of hind wing tubular. CUb vein
of hind wing not tubular. Hind wing with
3 hamuli. Metasoma: Median tergite of
first metasomal segment with pair of lat-
eral longitudinal carinae. First metasomal
median tergite without depression poster-
ad spiracle. Length width ratio of first me-
tasomal median tergite 0.98. Ovipositor
5.5-6.5 mm (6.3). Color: Head yellowish
orange. Antenna melanic. Maxillary and

labial palpomeres yellowish orange. Me-
sosoma yellowish orange. Forelegs yel-
lowish orange except femur and tibia me-
lanic, or melanic with yellowish orange
apically. Midlegs yellowish orange except
tibia melanic apically. Hindleg yellowish
orange except tibia yellowish orange with
apical third melanic and tarsus melanic
with basitarsus yellowish orange in basal
third. Forewing yellow basally and infus-
cate apically. Stigma melanic. Hind wing
yellow basally infuscate apically. Metaso-
ma yellowish orange. Ovipositor yellow-
ish orange.

6 . — Unknown.

Material examined. — Holotype: Costa
Rica: 9, Prov. Puntarenas, Est. Agujas, Rio
Agujas, sendero Samia, 300m, 1-3. vi. 1997,
A. Azofeifa (INBio). Paratypes: Costa Rica:
9, Rancho Quemado, 200m, Peninsula de
Osa, Prov. Puntarenas, vi.1992, F. Quesa-
da y M. Segura (INBio); 9, Rancho Que-
mado, Peninsula de Osa, Prov. Puntaren-
as, 200m, 01.xi-01.xii.1992, A. L. Marin
(INBio). Panama: 9, Barro Colorado Is,
9°9'N 79°51'W, 11-18.V.1994, J. Pickering
(UK).

Distribution. — This species is known
only from lowlands of Prov. Puntarenas,
Costa Rica and Barro Colorado in Panama.

Etymology. — From Greek bios meaning
life and pleres that means abundance.

Sesioctonus boliviensis Briceno, sp. n.

Diagnosis. — S. boliviensis can be separat-
ed by the following combination of char-
acters: interantennal space with a longi-
tudinal rounded keel and the presence of
the occipital tubercles.

Description. — 9. Length. Body, 10 mm.
Head: Interantennal space with rounded
longitudinal keel. Antennal sockets mod-
erately excavated. Face without median
longitudinal carina. Genae not expanded
posteroventrally. Occipital tubercles pre-
sent. Occiput excavated. Mandible con-
cave, outer tooth of mandible not longer
than inner tooth. Third and fourth labial
palpomeres completely fused. Mesosoma:

254

Journal of Hymenoptera Research

Subpronope triangular. Longitudinal ca-
rinae of scutellar depression absent. Scu-
tellum convex. Median areola of metano-
tum smooth; without median longitudinal
carina; and with lateral carinae present
and not meeting posteriorly. Propodeum
flat, median longitudinal carina absent.
Epicnemial carina sharp, complete, bi-
lobed medially (between forecoxae). Fore-
tibial spines absent. Midtibia with 6
spines. Hind tibia with 19 spines. Hind fe-
mur 3.21 times as long as wide. (RS + M)a
vein of forewing complete. 3RSa vein of
forewing absent. 2-1A vein of hind wing
tubular. CUb vein of hind wing not tu-
bular. Hind wing with 5 hamuli. Metaso-
ma: Median tergite of first metasomal seg-
ment without pair of lateral longitudinal
carinae. First metasomal median tergite
without depression posterad spiracle.
Length width ratio of first metasomal me-
dian tergite 0.79. Ovipositor 10 mm. Col-
or: Head melanic except labial palpomeres
melanic. Antenna melanic. Mesosoma yel-
lowish orange sometimes pronotum and
propleuron melanic. Forelegs melanic.
Midlegs melanic. Hindleg melanic. Fore-
wing banded from base infuscate, yellow,
infuscate. Stigma melanic. Hind wing en-
tirely infuscate. Metasoma yellowish or-
ange. Ovipositor yellowish orange.

S . — Unknown.

Material examined. — Holotype. Bolivia:
9, Staudinger K (ZSBS).

Distribution. — This species is known
only from Bolivia, South America.

Etymology. — The name boliviensis refers
to the country of origin of the holotype.

Sesioctomts brasiliensis Briceno, sp. n.

Figs. 14, 22, 27, 34

Diagnosis. — S. brasiliensis is the only spe-
cies with metasoma reddish brown and
forewing with a large hyaline spot. Also,
the median areola of metanotum does not
have longitudinal and lateral carinae, nor
longitudinal rugosities. Occipital tubercles
are present in this species.

Description.— 9 . Length. Body, 9-10 mm

(9.0). Head: Flagellum with 30-35 flagel-
lomeres (33). Interantennal space with
rounded longitudinal keel. Antennal sock-
ets deeply excavated, sometimes moder-
ately excavated. Face without median lon-
gitudinal carina. Genae moderately ex-
panded posteroventrally. Occipital tuber-
cles present. Occiput excavated. Mandible
concave, outer tooth of mandible not lon-
ger than inner tooth. Maxilla with 4 pal-
pomeres. Third and fourth labial palpo-
meres completely fused. Mesosoma: Sub-
pronope triangular. Longitudinal carinae
of scutellar depression absent. Scutellum
convex. Median areola of metanotum
smooth; without median longitudinal ca-
rina; and lacking lateral carinae. Propo-
deum convex, median longitudinal carina
absent. Epicnemial carina sharp, complete,
bilobed medially (between forecoxae).
Foretibial spines absent. Midtibia with 8-
11 spines (8). Hind tibia with 14-17 spines
(14). Hind femur 3.64 times as long as
wide. (RS+M)a vein of forewing com-
plete. 3RSa vein of forewing present. 2-1A
vein of hind wing tubular. CUb vein of
hind wing not tubular. Hind wing with 5-
6 hamuli (6). Metasoma: Median tergite of
first metasomal segment without pair of
lateral longitudinal carinae. First metaso-
mal median tergite without depression
posterad spiracle. Length width ratio of
first metasomal median tergite 1.09. Ovi-
positor 8 mm. Color: Head melanic. An-
tenna melanic. Maxillary palpomeres yel-
lowish orange. Labial palpomeres melan-
ic. Mesosoma melanic. Forelegs mostly
yellowish orange except coxae melanic,
tibia yellowish orange, or melanic; and
tarsus mostly yellowish orange, but apical
tarsomere melanic. Midlegs mostly melan-
ic except trochanter and trocantellus yel-
lowish orange, femur yellowish orange in
basal half, melanic apically. Hindlegs
mostly melanic except trochantellus me-
lanic, or yellowish orange. Forewing in-
fuscate with large hyaline spot. Stigma
melanic. Hind wing entirely infuscate.
Metasoma reddish brown with the last

Volume 12, Number 2, 2003

255

Figs. 28-32. Median areola of metanotum. 28, Smooth with lateral carinae not meeting posteriorly in S. diazi.
29, With longitudinal rugosities and lacking lateral carinae in S. grandis. 30-32, 30, Foretibia of S. acrolophus
showing spines. 31, Midtibia of S. dichromus showing spines. 32, Simple tarsal claws in S. acrolophus.

four segments melanic. Ovipositor yellow-
ish orange.

6 . — Essentially as the female.

Material examined. — Holotype. Brazil: 9,
Nova Teutonia, 27°11'S 52°23'W, 300-
500m, 24.L1939, Fritz Plaumann (CNC).
Paratypes. Brazil: 9, Nova Teutonia,

27°11'S 52°23'W, 300-500m, Fritz Plau-
mann; 29, same data except 30. i. 1939,
26.U939 (BMNH); 9, same data except
vii.1940; 6, same data except 18.V.1954
(AEI); 59, same data except ii.1967,
iii.1965, xi.1968, ii.1966; 6, same data ex-
cept 21.x. 1940 (CNC); 29, same data ex-

256 Journal of Hymenoptera Research

cept 6-10.iii.1967 (UK); 9, Represa Rio ange. Antenna melanic. Mesosoma mostly

Grande, Guanabara, xii.1967. M. Alvaren- yellowish orange except pronotum melan-

ga (AEI). ic anteriorly; metanotum, propodeum and

Distribution. — This species is known metapleuron yellowish orange or melanic;

only from Nova Teutonia region of Brazil, and propleuron always melanic. Forelegs

Etymology. — This species is named after with coxa, trochanter and trocantellus me-

the country of the holotype specimen. lanic, or yellowish orange; femur yellow-
ish orange, melanic basally; tibia and tar-

Sesioctonus chaconi Briceno, sp. n. sus yenowish orange. Midlegs with coxa

Diagnosis. — This species may be recog- yellowish orange ventrally, melanic dor-

nized by the combination of a long outer sally; trochanter melanic, trochantellus

tooth of the mandible and the absent of yellowish orange, femur melanic in basal

the 3RSa vein in the forewing. half, yellowish orange apically, or yellow-

Description. — 9. Length. Body, 6-9 mm ish orange, tibia yellowish orange, or me-

(6.5). Head: Flagellum with 29-33 flagel- lanic, and tarsus yellowish orange but api-

lomeres (29). Interantennal space with cal tarsomere melanic. Hindleg melanic

rounded longitudinal keel. Antennal sock- except coxa yellowish orange in basal half,

ets moderately excavated, or not excavat- melanic apically; femur melanic, or yel-

ed. Face without median longitudinal ca- lowish orange, melanic basally. Forewing

rina. Genae not expanded posteroventral- banded from base, yellow, infuscate, yel-

ly. Occipital tubercles present. Occiput ex- low, infuscate. Stigma melanic. Hind wing

cavated. Mandible concave, outer tooth of banded from base, yellow, infuscate, yel-

mandible longer than inner tooth. Maxilla low, infuscate. Metasoma yellowish or-

with 4 palpomeres. Third and fourth labial ange except first metasomal tergum yel-

palpomeres completely fused. Mesosoma: lowish orange, but median tergite melanic

Subpronope triangular. Longitudinal ca- centrally and fifth to eighth metasomal

rinae of scutellar depression absent. Scu- terga melanic. Ovipositor yellowish or-

tellum convex. Median areola of metano- ange.

turn smooth; without median longitudinal 6 . Essentially as female,

carina; and with lateral carinae present Material examined. — Holotype. Ecuador:

and not meeting posteriorly. Propodeum 9, Napo, Tena, 23.V.1977, DL& SS Vincent

flat, median longitudinal carina absent. (CNC). Paratypes. Ecuador: 9, Banos,

Epicnemial carina, sharp, complete, bi- 700m, 20.iii.1939, W. Clarke-Maclntyre

lobed medially (between forecoxae). Fore- (AEI); 9, Coca, v. 1965, Luis Pena (AEI); 9,

tibial spines absent. Midtibia with 8-13 Prov. Santa Clara, 30.vi.1976, P. M. Turner

spines (9). Hind tibia with 13-16 spines (CNC). Peru: 6, Yahuarmayo, 8.ii.l910,

(15). Hind femur 3.39 times as long as CHTTownsend col (CNC); 9, Loreto, Pu-

wide. (RS + M)a vein of forewing com- callpa, 12.vi.1951, J. M. Schuncke (CNC);

plete. 3RSa vein of forewing absent. 2-1 A 9, Loreto, Boqueron Abad, 27.xii.1961, J.

vein of hind wing tubular. CUb vein of M. Shuncke (CNC); 9, Avispas, 30m nr.

hind wing not tubular. Hind wing with 4- Marcapata, 1-15.X.1962, Luis Peha (AEI);

5 hamuli (4). Metasoma: Median tergite of 9, Loreto, Boqueron, 500m, 7-14.vii.1965,

first metasomal segment without pair of J. Schuncke (UK). Colombia: 9, Putuma-

lateral longitudinal carinae. First metaso- yo, Villa Garzon, 8mi. s. Mocoa,

mal median tergite without depression 3.viii.l978, M. Cooper (CNC).

postered spiracle. Length width ratio of Distribution.— -This species is distributed

first metasomal median tergite 1.19. Ovi- in northwestern of South America, from

positor 6-8 mm (6.1). Color: Head melanic Colombia to Peru and Ecuador.

vpt maxillary palpomeres yellowish or- Etymology.— This species is named in

Volume 12, Number 2, 2003

257

honor of Mr. Anibal Chacon, who during
many years has shared with Venezuelan
entomologists his passion for entomology
and insects collecting in Venezuela.

Sesioctonus chrestos Briceno, sp. n.

Diagnosis. — S. chrestos is known only
from one male specimen, however can be
distinguished from all other species by the
following combination of characters: max-
illa with 5 palpomeres, labium with 4, ep-
icnemial carina absent and foretibia with
spines.

Description. — 6. Length. Body, 5.5 mm.
Head: Flagellum with broken after flagel-
lomere 12. Interantennal space with
rounded longitudinal keel. Antennal sock-
ets not excavated. Face without median
longitudinal carina. Genae not expanded
posteroventrally. Occipital tubercles ab-
sent. Occiput not excavated. Maxilla with
5 palpomeres. Third and fourth labial pal-
pomeres partly fused. Mesosoma: Sub-
pronope oval. Longitudinal carinae of scu-
tellar depression absent. Scutellum con-
vex. Median areola of metanotum smooth;
without median longitudinal carina; and
with lateral carinae present and not meet-
ing posteriorly. Propodeum convex, me-
dian longitudinal carina absent. Epicne-
mial carina completely absent. Foretibial
spines present. Midtibia with 2 spines.
Hind tibia with 5 spines. Hind femur 3.06
times as long as wide. (RS + M)a vein of
forewing incomplete. 3RSa vein of fore-
wing present. 2-1 A vein of hind wing tu-
bular. CUb vein of hind wing not tubular.
Hind wing with 3 hamuli. Metasoma: Me-
dian tergite of first metasomal segment
without pair of lateral longitudinal cari-
nae. First metasomal median tergite with-
out depression posterad spiracle. Length
width ratio of first metasomal median ter-
gite 1.04. Color: Head yellowish orange.
Antenna melanic. Maxillary and labial
palpomeres yellowish orange. Mesosoma
yellowish orange except scutellum, meta-
notum, propodeum, mesopleuron and
metapleuron melanic. Forelegs yellowish

orange. Midlegs mostly melanic except fe-
mur yellowish orange apically; and tibia
yellowish orange. Hindleg melanic. Fore-
wing entirely infuscate. Stigma melanic.
Hind wing entirely infuscate. Metasoma
with first and second terga yellowish or-
ange, but median tergite melanic centrally;
rest of metasomal terga melanic.

9 . — Unknown.

Material examined. — Holotype. Peru: 6,
Marcapata (TMB).

Distribution. — This species is known
only from Peru.

Etymology. — From Greek chrestos that
means good, useful.

Sesioctonus clavijoi Briceno, sp. n.

Fig. 26

Diagnosis. — S. clavijoi shows color pat-
terns similar to S. areolatus, however can
be separated from the latter by the pres-
ence of the bilobed epicnemial carina. S.
areolatus has this carina straight and also
has a central areola on the propodeum
and a longitudinal carina on the scutellar
depression, both of which are absent in S.
clavijoi.

Description. — 9. Length. Body, 4-5.5 mm
(5.1). Head: Flagellum with 25 flagello-
meres. Interantennal space lacking longi-
tudinal keel. Antennal sockets not exca-
vated. Face without median longitudinal
carina. Genae not expanded posteroven-
trally. Occipital tubercles absent. Occiput
not excavated. Mandible concave, outer
tooth of mandible not longer than inner
tooth. Maxilla with 4 palpomeres. Third
and fourth labial palpomeres completely
fused. Mesosoma: Longitudinal carinae of
scutellar depression absent. Scutellum flat.
Median areola of metanotum smooth;
without median longitudinal carina; and
with lateral carinae present and meeting
posteriorly. Propodeum convex, median
longitudinal carina absent. Epicnemial ca-
rina sharp, complete, bilobed medially
(between forecoxae). Foretibial spines pre-
sent. Midtibia with 5-9 spines (8). Hind
tibia with 10-12 spines (12). Hind femur

258

Journal of Hymenoptera Research

3.1 times as long as wide. (RS+M)a vein
of forewing incomplete. 3RSa vein of fore-
wing absent. 2-1 A vein of hind wing not
tubular. CUb vein of hind wing not tu-
bular. Hind wing with 3 hamuli. Metaso-
ma: Median tergite of first metasomal seg-
ment with pair of lateral longitudinal ca-
rinae. First metasomal median tergite
without depression posterad spiracle.
Length width ratio of first metasomal me-
dian tergite 1.04. Ovipositor 4 mm. Color:
Head yellowish orange. Antenna melanic.
Maxillary and labial palpomeres yellow-
ish orange. Mesosoma yellowish orange.
Forelegs yellowish orange. Midlegs yel-
lowish orange. Hindleg yellowish orange
except tibia mostly yellowish orange, but
melanic apically and tarsus melanic, or
melanic with basitarsus yellowish orange
basally, or melanic with basitarsus yellow-
ish orange in basal third. Forewing entire-
ly infuscate. Stigma melanic. Hind wing
entirely infuscate. Metasoma yellowish or-
ange. Ovipositor yellowish orange.

6 . — Essentially as female.

Material examined. — Holotype. Costa
Rica: 9, Puntarenas, Golfo Dulce, 24km
W. Piedras Blancas, 200m, iv.1993, Paul
Hanson (UWY). Paratypes. Costa Rica: 39,
same data as holotype, except 10-200m,
xii.1992, ii.1993, Paul Hanson (UWY); 6,
Puntarenas, San Vito, Est. Biol. Las Altu-
ras, 1500m, iii.1992, Paul Hanson (UWY).
Peru: 9, Quincemil, 750m nr. Marcapata,
10-15.xi.1962, Luis Pena (AEI). Mexico: 9,
Vista Hermosa, Oaxaca, 96.5km SW Tux-
tepec, 19.X.1962, H. & M. Townes (AEI).

Distribution. — Southern Mexico, Costa
Rica and Peru.

Etymology. — This species is named in
honor of Jose Clavijo A., Venezuelan tax-
onomist, my professor and my friend,
who has shared with me many of my en-
tomologist dreams and has been an im-
portant part of my life. Thanks for all that
you have done for me.

Sesioctonus diazi Briceno, sp. n.

Figs. 17, 28

—S. diazi is not an easy spe-
distinguish because of intraspecific

variation, especially in color pattern. It can
be confused with specimens of S. chaconi,
and separation of these species is compli-
cated because the differences are primarily
in the relative sizes of the mandible teeth.
Description. — 9. Length. Body, 7-8 mm
(7.5). Head: Flagellum with 28-34 flagel-
lomeres (33). Interantennal space with
rounded longitudinal keel. Antennal sock-
ets moderately excavated. Face without
median longitudinal carina. Genae not ex-
panded posteroventrally. Occipital tuber-
cles present. Occiput excavated. Mandible
concave, outer tooth of mandible not lon-
ger than inner tooth. Maxilla with 4 pal-
pomeres. Third and fourth labial palpo-
meres completely fused. Mesosoma: Sub-
pronope triangular or oval-shape. Longi-
tudinal carinae of scutellar depression
absent. Scutellum convex. Median areola
of metanotum smooth; without median
longitudinal carina; and with lateral cari-
nae present and not meeting posteriorly.
Propodeum flat, median longitudinal ca-
rina absent. Epicnemial carina, sharp,
complete, bilobed medially (between fore-
coxae). Foretibial spines absent. Midtibia
with 8-11 spines (10). Hind tibia with 15-
23 spines (17). Hind femur 3.01-3.22 (3.08)
times as long as wide. (RS + M)a vein of
forewing complete or incomplete. 3RSa
vein of forewing present. 2-1 A vein of
hind wing tubular. CUb vein of hind wing
not tubular. Hind wing with 4-5 (5) ham-
uli. Metasoma: Median tergite of first me-
tasomal segment without pair of lateral
longitudinal carinae. First metasomal me-
dian tergite without depression posterad
spiracle. Length width ratio of first meta-
somal median tergite 0.84-1.03 (0.84). Ovi-
positor 5-8 mm (7.1). Color: Head melan-
ic. Antenna melanic. Maxillary and labial
palpomeres yellowish orange except basal
two palpomeres melanic. Mesosoma most-
ly yellowish orange sometimes pronotum
and propleuron melanic. Forelegs melanic,
or tarsus mostly yellowish orange, but
apical tarsomere melanic. Midlegs vari-
able, tarsus melanic, or yellowish orange

Volume 12, Number 2, 2003

259

Figs. 33-37. First metasomal tergite. 33,35, With lateral longitudinal carinae. 33, S. dichromus. 35, S. acrolophus.
34, S. brasiliensis without lateral longitudinal carinae. 36-37, S. grandis showing depression posterad spiracle.

but apical tarsomere melanic. Hindlegs
melanic but coxa can be yellowish orange
in basal half, melanic apically, or yellow-
ish orange with melanic spots laterally, or
melanic, or yellowish orange basally, oth-
erwise melanic. Forewing yellow basally
and infuscate apically, or banded from

base, yellow, infuscate, yellow, infuscate.
Stigma melanic. Hind wing yellow basally
infuscate apically. Metasoma yellowish or-
ange. Ovipositor yellowish orange, or yel-
lowish orange except apical eighth melan-
ic.

6 . — The color pattern in males is essen-

260

Journal of Hymenoptera Research

tially as females except fifth to eighth me-
tasomal tergite which sometimes can be
yellowish orange with central portion of
median tergite melanic.

Material examined. — Holotype. Costa
Rica: 9, Heredia, Est. Biol. La Selva, 50-
150m, 10°26'N 84°01'W, viii.1998 (INBio).
Paratypes. Costa Rica: 9, Nara NE Quepos,
16.vii.1975, W.J.Hanson (CNC); 9, Prov.
Alajuela, 28.V.1972, J.F.Alvarez (UCR); 6,
Alajuela, Upala, Colonia Libertad, 450m,
1-6. v. 1988, Gonzalez & Soto (UCR); 9,
Prov. Golfo Dulce, 24km W Pan-Am high-
way, 200m, iii-v.1989, Gauld (BMNH);
39, 2c?, same data as holotype except,
02.V.1993, vi.1993, vii.1993, 03.viii.1993,
v.1996 (INBio)(UCOB)(UK); 9, Prov. Pun-
tarenas, Est. Agujas, sendero Ajo., 300m,
14-24.viii.1996, A. Azofeifa (INBio). Gua-
temala: 9, Concepcion, 1400m (UK). Pan-
ama: 49, Barro Colorado Is, 9°9'N
79°51'W, v.1939, Jas Zetek (USNM), 5-
12.V.1993, 23-30.iii.1994, 30.iii-6.iv.1994. J.
Pickering (UK); 19, 16, Portobello,
24.ii.1911, 13.iii.1911, A. Busk (CNC); 9,
San Bias Nusagandi Reserve, 9°20'N
79°0'W, 20-27.xi.1993. J. Pickering (UK).

Distribution. — Sesioctonus diazi is distrib-
uted from Guatemala to Panama in Cen-
tral America.

Etymology. — This species is named in
honor of Francisco Diaz, Venezuelan en-
tomologist, for his contribution to the
knowledge of Venezuela ichneumonids.

Sesioctonus dichromus Briceno, sp. n.

Figs. 12, 16, 18, 20, 31, 33

Diagnosis. — S. dichromus can be distin-
guished from other Sesioctonus species by
the following combination of characters:
occipital tubercles present, occiput exca-
vated, median areola of metanotum with
lateral carinae present and meeting pos-
teriorly, the median tergite of first meta-
somal segment with well defined pair of
lateral longitudinal carinae.

Description.— 9. Length. Body, 6-10 mm
(9.5). Head: Flagellum with 30-35 flagel-
lomeres (33). Interantennal space with

rounded longitudinal keel. Antennal sock-
ets moderately excavated. Face without
median longitudinal carina. Genae not ex-
panded posteroventrally. Occipital tuber-
cles present. Occiput excavated. Mandible
concave, outer tooth of mandible longer
than inner tooth. Maxilla with 4 palpo-
meres. Third and fourth labial palpomeres
completely fused. Mesosoma: Subpronope
triangular or oval. Longitudinal carinae of
scutellar depression absent. Scutellum
convex. Median areola of metanotum
smooth; without median longitudinal ca-
rina; and with lateral carinae present and
meeting posteriorly, or lacking lateral ca-
rinae. Propodeum flat, median longitudi-
nal carina present or absent. Epicnemial
carina sharp, complete, bilobed medially
(between forecoxae). Foretibia spines ab-
sent. Midtibia with 7-13 spines (12). Hind
tibia with 18-25 spines (19). Hind femur
3.28-3.6 times as long as wide (3.54).
(RS+M)a vein of forewing complete or in-
complete. 3RSa vein of forewing present
or absent. 2-1 A vein of hind wing tubular.
CUb vein of hind wing not tubular. Hind
wing with 5-7 hamuli (7). Metasoma: Me-
dian tergite of first metasomal segment
with pair of lateral longitudinal carinae.
Median tergite of first metasomal segment
without depression posterad spiracle.
Length width ratio of first metasomal me-
dian tergite 0.97. Ovipositor 6-9 mm (9.0).
Color: Head melanic including maxillary
and labial palpomeres. Antenna melanic.
Mesosoma melanic. Forelegs melanic.
Midlegs melanic. Hindleg melanic. Fore-
wing entirely infuscate, or hyaline basally,
infuscate apically. Stigma melanic. Hind
wing entirely infuscate, or infuscate with
large hyaline spots, or hyaline basally, in-
fuscate apically. Metasoma yellowish or-
ange. Ovipositor yellowish orange.

6 . — Essentially as the female.

Material examined. — Holotype. Costa
Rica: 9, Alajuela, San Ramon/800m, 29.i-
03.ii.1995, G. Carballo (INBio). Paratypes.
Costa Rica: 29 9, Heredia, Est. Biol. La Sel-
va, 50-150m, 10°26'N 84°01'W, 11-

Volume 12, Number 2, 2003

261

17. vi. 1986, 2. v. 1993, vi.1993, viii.1993,
ix.1993, 01. ix. 1993, 16. ix. 1993, xii.1993,
22.iii.1994, 29.ix.1995, ix.1995, 15.xii.1995,
14.xii.1995, ii.1996, 01.iii.1996, iii.1996,
vi.1996, 2. v. 1996, 15. v. 1996, 02.x. 1997,
xi.1997, ix.1998, 22.L1999 (INBio) (UK)
(UCR) (UWY) (UCOB); 6, Heredia, Est. El
Ceibo, Braulio Carrillo, N.P. 400-600m,
iii.1990, C. Chavez (INBio); 29, Est. Pitilla,
9km Sur Santa Cecilia, Guanacaste, 700m,
xi.1988, C. Chavez & M. Espinoza, ii.1990,
P. Rios, C. Moraga & R. Blanco (INBio);
4 9, El Limon, Sector Cerro Cocori, Finca
de E. Rojas, 150m, v.1991, . 5.vii-12.viii.
1992, ll.i-12.ii.1993, Trampa malaise (IN-
Bio); 9, El Limon, 16km W Guapiles,
Parque Nacional Braulio Carrillo, 400m,
iv-v.1989, Gauld (BMHN); 9, Limon, P N
Tortuguero, Est. 4-esquinas, 0m, vi-
viii.1989, Solano col (UWY); 9, Puntaren-
as, Rancho Quemado, Peninsula de Osa,
200m, iv.1992, L.Brenes (INBio); 9, Ala-
juela, Penas Blancas, 700m, viii.1987, E.
Cruz (CNC); 9, Alajuela, Sector Colonia
Palmareha, 9km SO de Bajo Rodriguez,
700m, ix.1996, G. Carballo (INBio); 9, San
Jose, P.N. Braulio Carrillo, 9.5km E tunel,
1000m, viii-ix.1989 (UWY); 6\ Alajuela,
Sector San Ramon, 800m, ll-15.iv.1994,
M. Zumbado (INBio); 9, Est. Biol. La Sel-
va, 30.vi.1995 (UK). Mexico: 9, Chiapas,
Muste, 440m near Huixtla. 1970. Mai. trap.
Welling (CNC).

Distribution. — This species is known
from Mexico to Costa Rica.

Etymology. — From Greek di that means
two and chromos that means color, in ref-
erence to the color pattern of the species,
half melanic, half yellowish orange.

Sesioctomis dominions Briceno, sp. n.

Diagnosis. — S. dominicus is known only
from males specimens. However, it can be
separated from all other Sesioctomis spe-
cies by its small size (2.0-3. 5mm) and to-
tally hyaline wings.

Description. — 6 . Length. Body, 2-3.5 mm
(2.5). Head: Flagellum with 23-27 flagel-
lomeres (25). Interantennal space lacking

longitudinal keel. Antennal sockets not ex-
cavated. Face without median longitudi-
nal carina. Genae not expanded poster-
oventrally. Occipital tubercles absent. Oc-
ciput not excavated. Mandible flat, outer
tooth of mandible not longer than inner
tooth. Maxilla with 4 palpomeres. Third
and fourth labial palpomeres completely
fused. Mesosoma: Subpronope elongate-
oval. Longitudinal carinae of scutellar de-
pression absent. Scutellum convex. Medi-
an areola of metanotum smooth; without
median longitudinal carina; and with lat-
eral carinae present and meeting posteri-
orly. Propodeum convex, median longi-
tudinal carina absent. Epicnemial carina
(difficult to see), sharp, incomplete dorsal-
ly, straight medially (between forecoxae).
Foretibial spines absent. Midtibia with 7-
10 spines (8). Hind tibia with 13-16 spines
(15). Hind femur 1.14 times as long as
wide. (RS + M)a vein of forewing incom-
plete. 3RSa vein of forewing absent. 2-1A
vein of hind wing not tubular. CUb vein
of hind wing not tubular. Hind wing with
3 hamuli. Metasoma: Median tergite of
first metasomal segment without pair of
lateral longitudinal carinae. First metaso-
mal median tergite without depression
posterad spiracle. Length width ratio of
first metasomal median tergite 1.10. Color:
Head black except face yellowish orange.
Antenna melanic. Maxillary and labial
palpomeres yellowish orange. Mesosoma
melanic except propodeum, propleuron
and metapleuron yellowish orange and
pronotum yellowish orange, or melanic.
Forelegs yellowish orange. Midlegs yel-
lowish orange except tibia melanic, or yel-
lowish orange. Hindleg yellowish orange
except tibia melanic and tarsus melanic.
Forewing hyaline. Stigma melanic. Hind
wing hyaline. Metasoma with first and
second terga yellowish orange, the rest
melanic.

9 . — Unknown.

Material examined. — Holotype. Dominica
W.I.: 6, Springfield, xi.1967, N.L.H Krauss
(UK). Paratypes. Dominica W.I: 2<5, Clarke

262

Journal of Hymenoptera Research

Hall, Layou Vail, 10-17.ii.1965, 20-
28.ii.1965, H.E.Evans (UK) (USNM); 6\
Mth. Layou R, 13.iii.1965, H.E.Evans
(USNM); 6, Hillsborough, 15.iii.1965
(USNM).

Distribution. — This species is known
only from the Dominica W. I. in the Ca-
ribbean.

Etymology. — This species is named after
the locality of the holotype specimen.

Sesioctomis enmenetes Briceno, sp. n.

Diagnosis. — S. enmenetes is known only
from one specimen, however, can be dis-
tinguised from all other Sesioctomis species
by the following combination of charac-
ters: occipital tubercles absent, presence of
median longitudinal carina on the propo-
deum, epicnemial carina complete, medi-
an areola of metanotum smooth and a pair
of longitudinal carinae on the first meta-
somal tergite.

Description. — 9. Length. Body, 6 mm.
Head: Flagellum broken after flagellomere
11. Interantennal space with rounded lon-
gitudinal keel. Antennal sockets not exca-
vated. Face without median longitudinal
carina. Genae not expanded posteroven-
trally. Occipital tubercles absent. Occiput
excavated. Mandible concave, outer tooth
of mandible longer than inner tooth. Max-
illa with 4 palpomeres. Third and fourth
labial palpomeres completely fused. Me-
sosoma: Subpronope triangular. Longitu-
dinal carinae of scutellar depression ab-
sent. Scutellum convex. Median areola of
metanotum smooth; without median lon-
gitudinal carina; and with lateral carinae
present and not meeting posteriorly. Pro-
podeum flat, median longitudinal carina
absent. Epicnemial carina sharp, complete,
bilobed medially (between forecoxae).
Foretibia spines absent. Midtibia with 10
spines. Hind tibia with 15 spines. Hind fe-
mur 3.17 times as long as wide. (RS+M)a
vein of forewing incomplete. 3RSa vein of
forewing present. 2-1A vein of hind wing
eir. CUb vein of hind wing not fu-
nd wing with 5 hamuli. Metaso-

ma: Median tergite of first metasomal seg-
ment with pair of lateral longitudinal ca-
rinae. First metasomal median tergite
without depression posterad spiracle.
Length width ratio of first metasomal me-
dian tergite 1.09. Ovipositor 6.5 mm. Col-
or: Head melanic. Antenna melanic. Max-
illary and labial palpomeres melanic. Me-
sosoma yellowish orange except prono-
tum mostly yellowish orange, melanic
anteriorly and propleuron melanic. Fore-
legs melanic except coxa mostly yellowish
orange and tarsus mostly yellowish or-
ange, but apical tarsomere melanic. Mid-
legs melanic except coxa yellowish or-
ange. Hindleg melanic except coxa yel-
lowish orange and femur melanic in basal
third, otherwise yellowish orange. Fore-
wing entirely infuscate. Stigma melanic.
Hind wing entirely infuscate. Metasoma
yellowish orange. Ovipositor yellowish
orange except apical eighth melanic.

(5 . — Unknown.

Material examined. — Holotype. Costa
Rica: 9, San Vito Las Cruces, 1200m, 9.vii-
7.viii.l982, B. Gill (CNC).

Distribution. — This species is known
only from the Las Cruces region of Costa
Rica.

Etymology. — From Greek enmenetes that
means friend, in honor of my colleagues
and friends Ana, Carmen Liceth, Carlos,
Dorys and Maria del Carmen.

Sesioctomis galeos Briceno, sp. n.

Diagnosis. — S. galeos shows the follow-
ing combination of characters: occipital tu-
bercles absent, occiput not excavated,
maxillary palpi with 4 palpomeres and la-
bial palpi with 3, median areola of meta-
notum with longitudinal rugosities, epic-
nemial carina incomplete laterally and
straight medially.

Description. — 9. Length. Body, 8.5 mm.
Head: Flagellum with 33 flagellomeres. In-
terantennal space with rounded longitu-
dinal keel, or lacking of longitudinal keel.
Antennal sockets deeply excavated, or
moderately excavated. Face without me-

Volume 12, Number 2, 2003

263

dian longitudinal carina. Genae moderate-
ly expanded posteroventrally. Occipital
tubercles absent. Occiput not excavated.
Mandible concave, outer tooth of mandi-
ble not longer than inner tooth. Maxilla
with 4 palpomeres. Third and fourth labial
palpomeres completely fused. Mesosoma:
Subpronope triangular. Longitudinal ca-
rinae of scutellar depression absent. Scu-
tellum convex. Median areola of metano-
tum with longitudinal rugosities; without
median longitudinal carina; and with lat-
eral carinae present and not meeting pos-
teriorly. Propodeum flat, median longitu-
dinal carina of propodeum absent. Epic-
nemial carina sharp, incomplete laterally,
straight medially (between forecoxae).
Foretibia spines absent. Midtibia with 2-7
(2) spines. Hind tibia with 8-12 (12)
spines. Hind femur 3.6 times as long as
wide. (RS + M)a vein of forewing com-
plete. 3RSa vein of forewing present or ab-
sent. 2-1 A vein of hind wing tubular or
not tubular. CUb vein of hind wing not
tubular. Hind wing with 4-5 (5) hamuli.
Metasonta: Median tergite of first meta-
somal segment without pair of lateral lon-
gitudinal carinae. First metasomal median
tergite without depression posterad spi-
racle. Length width ratio of first metaso-
mal median tergite 0.9. Ovipositor 7.2mm.
Color: Head melanic. Antenna melanic.
Maxillary and labial palpomeres yellow-
ish orange. Mesosoma with pronotum yel-
lowish orange, sometimes melanic anteri-
orly; mesoscutum, scutellum and meta-
notum yellowish orange; propodeum
mostly yellowish orange with melanic
spots; propleuron yellowish orange or me-
lanic; mesopleuron mostly yellowish or-
ange, melanic basally and metapleuron
melanic, or yellowish orange. Forelegs yel-
lowish orange. Midlegs yellowish orange.
Hindleg melanic except trochanter and
trochantellus yellowish orange. Forewing
banded from base, yellow, infuscate, yel-
low, infuscate. Stigma melanic. Hind wing
banded from base, yellow, infuscate, yel-
low, infuscate. Metasoma mostly yellow-

ish orange but fourth tergum with median
tergum melanic and fifth to eighth terga
melanic.

8 . — Essentially as female.

Material examined. — Holotype. Brasil: 9,
Manaus, Reserva Dulce, 31.viii.1990,
Trampa malaise, Vidal col. (INPA). Para-
types. Peru: 6, Puerto Bermudez, 12-
19.vii.1920, Cornell Univ. Expedition
(CUIC); 9, Avispas, 30km nr Marcapata,
ix.1992, Luis Pena (AEI).

Distribution. — Sesioctonus galeos is
known only from Brazil and Peru.

Etymology. — From Greek galeos that
means shark in honor of Michael Sharkey,
in acknowledgment for his advising in my
formation as braconid taxonomist and for
his contribution to the knowledge of the
Agathidinae of the World.

Sesioctonus garciai Briceno, sp. n.

Diagnosis. — S. garciai is known only
from one specimen and can be distin-
guished from all other species by the fol-
lowing combination of characters: occipi-
tal tubercles present, occiput excavated,
epicnemial carina complete and straight
medially, although the presence of spines
on the fore tibia.

Description. — 9. Length. Body, excluding
ovipositor, 6 mm. Head: Flagellum with 30
flagellomeres. Interantennal space with
rounded longitudinal keel. Antennal sock-
ets not excavated. Face without median
longitudinal carina. Genae not expanded
posteroventrally. Occipital tubercles pre-
sent. Occiput excavated. Mandible con-
cave, outer tooth of mandible longer than
inner tooth. Maxilla with 4 palpomeres.
Third and fourth labial palpomeres com-
pletely fused. Mesosoma: Subpronope tri-
angular. Longitudinal carinae of scutellar
depression absent. Scutellum convex. Me-
dian areola of metanotum smooth; with-
out median longitudinal carina; and with
lateral carinae present and not meeting
posteriorly. Propodeum flat, median lon-
gitudinal carina of propodeum absent. Ep-
icnemial carina sharp, complete, straight

264

Journal of Hymenoptera Research

medially (between forecoxae). Foretibia
spines present. Midtibia with 9 spines.
Hind tibia with 15 spines. Hind femur 3.64
times as long as wide. (RS + M)a vein of
forewing incomplete. 3RSa vein of fore-
wing absent. 2-1 A vein of hind wing tu-
bular. CUb vein of hind wing not tubular.
Hind wing with 4 hamuli. Metasoma: Me-
dian tergite of first metasomal segment
without pair of lateral longitudinal cari-
nae. First metasomal median tergite with-
out depression posterad spiracle. Length
width ratio of first metasomal median ter-
gite 0.9. Ovipositor 5 mm. Color: Head
melanic. Antenna melanic. Maxillary and
labial palpomeres yellowish orange except
two basal palpomeres melanic. Mesosoma
yellowish orange. Forelegs yellowish or-
ange. Midlegs yellowish orange except
tibia and tarsus melanic. Hindleg melanic
with coxa yellowish orange femur mostly
yellowish orange, but melanic distally.
Forewing entirely infuscate. Stigma me-
lanic. Hind wing entirely infuscate. Meta-
soma tergum yellowish orange except fifth
to eighth metasomal terga mostly yellow-
ish orange but median tergites melanic
posteriorly. Ovipositor yellowish orange
except apical eighth melanic.

6 . — Unknown.

Materia] examined. — Holotype. Brasil: 9,
Nova Teutonia, 27°11'N 52°23'L, 2.ii.l939,
Fritz Plaumann (CNC).

Distribution. — This species is known
only from the Nova Teutonia region of
Brazil.

Etymology. — This species is named in
honor of Jose Luis Garcia, Venezuelan en-
tomologist, for his contribution to the
knowledge of Venezuela proctotrupoids.

Sesioctonus grandis Briceno, sp. n.

Figs. 19, 29, 36, 37

Diagnosis. — S. grandis is the larger spe-
cies of Genus Sesioctonus and it has a high-
er number of flagellomeres (48), maxilla
and labium with four palpomeres, and the
first metasomal median tergite with de-
pression posterad spiracle. This latter

character is shared with S. qui from which
it is separated by the presence of an oval-
shape subpronope, epicnemial carina ab-
sent and the presence of a pair of lateral
longitudinal carinae on the first metaso-
mal median tergite.

Description. — 9. Length. Body, excluding
ovipositor, 10-13 mm (10.0). Head: Flagel-
lum with 44-48 flagellomeres (46). Inter-
antennal space with sharp longitudinal
keel. Antennal sockets deeply excavated.
Face without median longitudinal carina.
Genae strongly expanded posteroventral-
ly. Occipital tubercles absent. Occiput not
excavated. Mandible flat, outer tooth of
mandible not longer than inner tooth.
Maxilla with 4 palpomeres. Third and
fourth labial palpomeres partly fused. Me-
sosoma: Subpronope oval. Longitudinal
carinae of scutellar depression absent.
Scutellum convex. Median areola of me-
tanotum with longitudinal rugosities;
without median longitudinal carina; and
lacking lateral carinae. Propodeum con-
vex, median longitudinal carina of pro-
podeum absent. Epicnemial carina com-
pletely absent. Foretibia spines absent.
Midtibia with 5-8 spines (8). Hind tibia
with 14-21 spines (21). Hind femur 4.4
times as long as wide. (RS + M)a vein of
forewing complete. 3RSa vein of forewing
absent. 2-1 A vein of hind wing not tubu-
lar. CUb vein of hind wing not tubular.
Hind wing with 8-10 hamuli (9). Meta-
soma: Median tergite of first metasomal
segment with pair of lateral longitudinal
carinae. First metasomal median tergite
with depression posterad spiracle. Length
width ratio of first metasomal median ter-
gite 0.71. Ovipositor 10 mm. Color: Head
melanic. Antenna melanic, sometimes yel-
lowish orange. Maxillary and labial pal-
pomeres yellowish orange except two bas-
al palpomeres melanic. Mesosoma melan-
ic. Forelegs yellowish orange except coxa
melanic, trochantellus, tibia and tarsus
yellowish orange, or melanic. Midlegs yel-
lowish orange except coxae and tarsus
melanic. Hindleg melanic. Forewing

Volume 12, Number 2, 2003

265

banded from base, yellow, infuscate, yel-
low, infuscate. Stigma melanic, or melanic
and yellowish orange. Hind wing yellow
basally infuscate apically. Metasoma me-
lanic. Ovipositor yellowish orange.

6 . — Essentially as the female.

Material examinated. — Holotype. Brazil:
9, Rio Grande do Sul, Staudinger K col.
(ZSBS). Paratypes. Brazil: 49, 56, Rio
Grande do Sul, Staudinger K col. (ZSBS)
(UCOB).

Distribution. — This species is known
only from the Rio Grande do Sul region
of Brazil.

Etymology. — grandis refers the big size of
these specimens.

Sesioctonus kompsos Briceno, sp. n.

Diagnosis. — S. kompsos can be distin-
guished for the following combination of
characters: occipital tubercles present, oc-
ciput excavated, median areola of meta-
notum with longitudinal rugosities.

Description. — 9. Length. Body, excluding
ovipositor, 8.0-10. 0mm (10). Head: Flagel-
lum with 33-34 (34) flagellomeres. Inter-
antennal space with rounded longitudinal
keel. Antennal sockets not excavated. Face
without median longitudinal carina. Ge-
rtae not expanded posteroventrally. Occip-
ital tubercles present. Occiput excavated.
Mandible concave, outer tooth of mandi-
ble longer than inner tooth. Maxilla with
4 palpomeres. Third and fourth labial pal-
pomeres completely fused. Mesosoma:
Subpronope triangular. Longitudinal ca-
rinae of scutellar depression absent. Scu-
tellum convex. Median areola of metano-
tum with longitudinal rugosities; without
median longitudinal carina; and with lat-
eral carinae present and not meeting pos-
teriorly. Propodeum flat, median longitu-
dinal carina absent. Epicnemial carina
sharp, complete, bilobed medially (be-
tween forecoxae). Foretibia spines absent.
Midtibia with 9-13 (13) spines. Hind tibia
with 21-25 (25) spines. Hind femur 3.2-
3.43 (3.2) times as long as wide. (RS + M)a
vein of forewing complete. 3RSa vein of

forewing absent. 2-1 A vein of hind wing
tubular. CUb vein of hind wing not tu-
bular. Hind wing with 6 hamuli. Metaso-
ma: Median tergite of first metasomal seg-
ment without pair of lateral longitudinal
carinae. First metasomal median tergite
without depression posterad spiracle.
Length width ratio of first metasomal me-
dian tergite 0.94-1.20 (0.94). Ovipositor 8
mm. Color: Head melanic. Antenna me-
lanic. Maxillary palpomeres yellowish or-
ange except two basal palpomeres melan-
ic. Labial palpomeres melanic. Mesosoma
mostly melanic except metanotum, pro-
podeum and metapleuron yellowish or-
ange. Forelegs melanic. Midlegs melanic.
Hindleg melanic. Forewing entirely infus-
cate. Stigma melanic. Hind wing entirely
infuscate. Metasoma yellowish orange.
Ovipositor yellowish orange except apical
eighth melanic.

6 . — Unknown.

Material examined. — Holotype. Costa
Rica: 9, Rancho Quemado, 2km N. cami-
no Drake, 275m, i.1991, P. Hanson (UCR).
Paratype. Brasil: 9, Guanabara, Represa
Rio Grande, viii.1966, M.Alvarenga (AEI).

Distribution. — This species is known
only from Costa Rica and Brazil.

Etymology. — From Greek kompsos that
means elegance in reference to the beauty
of this species.

Sesioctonus miyayensis Briceno, sp. n.

Diagnosis. — S. miyayensis shows the fol-
lowing combination of characters: occipi-
tal tubercles absent, subpronope elongate-
oval-shaped, median areola of metanotum
with lateral carinae meeting posteriorly,
epicnemial carinae complete and straight
medially, foretibia with spines, first me-
tasomal tergite with pair of lateral longi-
tudinal carinae.

Description.— 9 . Length. Body, excluding
ovipositor, 5-7 mm (6.5). Head: Flagellum
with 32 flagellomeres. Interantennal space
with rounded longitudinal keel. Antennal
sockets not excavated. Face without me-
dian longitudinal carina. Genae moderate-

Journal of Hymenoptera Research

ly expanded posteroventrally. Occipital (AEI). Paratopes. Costa Rica: 89, Heredia,

tubercles absent. Occiput not excavated. Est. Biol. La Selva, 10°26'N 84°01'W, 50-

Mandible flat, outer tooth of mandible not 150m, xii.1992, P. Hanson (UWY),

longer than inner tooth. Maxilla with 4 l.xi.1993, J. Longino (UK), 01.viii.1995,

palpomeres. Third and fourth labial pal- 15.1.1996, iii.1996, ii.1996, 31. v. 1996,

pomeres completely fused. Mesosoma: 19.ii.1998 (UK); 9, Prov. Guanacaste, Est.

Subpronope elongate-oval. Longitudinal Pitilla 9km S. Sta. Cecilia, 700m, 4-

carinae of scutellar depression absent. 14.xi.1991, D.Garcia (INBio); 9, Prov. Pun-

Scutellum convex. Median areola of me- tarenas, Rancho Quemado, Peninsula de

tanotum smooth; without median longi- Osa, 200m, vi.1992. F.Quesada y M.Segura

tudinal carina; and with lateral carinae (INBio).

present and meeting posteriorly. Propo- Distribution.— This species is known

deum flat, median longitudinal carina ab- only from Costa Rica,

sent. Epicnemial carina sharp, complete, Etymology.— This species is named in

straight medially (between forecoxae). honor of my father's birthplace, Miyayi.
Foretibia spines present. Midtibia with 5-

7 spines (7). Hind tibia with 14 spines. Sesioctonus parathyridis Viereck

Hind femur 3.42 times as long as wide. ^^ tln/ndis Viereck 1912: h 9> »Par.

(RS + M)a vein of forewing incomplete. ^ Canal de panama„ (Washington/ 9#

3RSa vein of forewing absent. 2-1A vein i4552)._Busk 1912: 10, fig. (host a). Host a:

of hind wing tubular. CUb vein of hind Parathyridis perspicilla Stoll.
wing not tubular. Hind wing with 3-4

hamuli (3). Metasoma: Median tergite of Diagnosis. — S. parathyridis shares char-
first metasomal segment with pair of lat- acters with S. grandis and S. qui. They con-
eral longitudinal carinae. First metasomal form the group of species with the larger
median tergite without depression poster- size and the number of flagellomeres more
ad spiracle. Length width ratio of first me- than 40. Also, they have the interantennal
tasomal median tergite 1.1. Ovipositor 5 space with a longitudinal sharp keel,
mm. Color: Head yellowish orange. An- which is present in S. acrolophus. However,
tenna melanic. Maxillary and labial pal- S. parathyridis can be separated from these
pomeres melanic, or yellowish orange ex- species by the presence of a pair of lateral
cept two basal palpomeres melanic. Me- longitudinal carinae on the median tergite
sosoma yellowish orange. Forelegs yel- of first metasomal segment. This character
lowish orange except femur and tibia is shared with S. grandis. However, this
melanic. Midlegs yellowish orange except latter species has the first metasomal seg-
tibia yellowish orange basally, otherwise ment with a depression posterad spiracle,
melanic and tarsus melanic with basitar- which are absent in parathyridis.
sus yellowish orange in basal half. Hind- Description. — 9. Length. Body, excluding
leg yellowish orange except tibia yellow- ovipositor, 9-12 mm (12.0). Head: Flagel-
ish orange in basal half, melanic apically lum with 44-45 flagellomeres (42). Inter-
and tarsus melanic. Forewing yellow ba- antennal space with sharp longitudinal
sally and infuscate apically. Stigma melan- keel. Antennal sockets deeply excavated,
ic, or yellowish orange. Hind wing yellow Face without median longitudinal carina,
basally infuscate apically. Metasoma yel- Genae strongly expanded posteroventral-
lowish orange. Ovipositor yellowish or- ly. Occipital tubercles absent. Occiput not
ange. excavated. Mandible flat, outer tooth not
(J— Unknown. longer than inner tooth. Maxilla with 4
Material examined.— Holotype. Costa palpomeres. Third and fourth labial pal-
Rica: 9, Pto. Viejo, 50m, ii. 1980, W. Mason pomeres completely fused. Mesosoma:

Volume 12, Number 2, 2003

267

Subpronope triangular. Longitudinal ca-
rinae of scutellar depression absent. Scu-
tellum convex. Median areola of metano-
tum with longitudinal rugosities; without
median longitudinal carina; and lacking
lateral carinae. Propodeum flat, median
longitudinal carina absent. Epicnemial ca-
rina blunt, incomplete laterally, bilobed
medially (between forecoxae). Foretibia
spines absent. Midtibia with 5-8 spines
(6). Hind tibia with 14-16 spines (14).
Hind femur 4.88 times as long as wide.
(RS + M)a vein of forewing complete. 3RSa
vein of forewing present. 2-1A vein of
hind wing not tubular. CUb vein of hind
wing not tubular. Hind wing with 7-8
hamuli (7). Metasoma: Median tergite of
first metasomal segment with pair of lat-
eral longitudinal carinae, or without pair
of lateral longitudinal carinae. First meta-
somal median tergite without depression
posterad spiracle. Length width ratio of
first metasomal median tergite 0.8. Ovi-
positor 10-12 mm (12.0). Color: Head me-
lanic. Antenna melanic. Maxillary and la-
bial palpomeres yellowish orange except
two basal palpomeres melanic. Pronotum
mostly melanic with yellowish orange ar-
eas. Mesoscutum yellowish orange, or
mostly melanic, yellowish orange dorsal-
ly. Scutellum yellowish orange. Metano-
tum yellowish orange. Propodeum yel-
lowish orange, or mostly yellowish orange
with melanic spots. Propleuron mostly
melanic with yellowish orange areas, or
yellowish orange. Mesopleuron yellowish
orange. Metapleuron yellowish orange.
Forelegs mostly yellowish orange except
femur melanic; tibia yellowish orange, or
melanic but yellowish orange distally and
tarsus mostly yellowish orange, but apical
tarsomere melanic. Midlegs mostly yel-
lowish orange with variations on coxa
sometimes melanic apically; femur yel-
lowish orange in basal half, melanic api-
cally; tibia yellowish orange in basal half,
melanic apically, or yellowish orange and
tarsus melanic. Hindleg melanic except
coxa yellowish orange in basal half, me-

lanic apically. Forewing banded from
base, yellow, infuscate, yellow, infuscate.
Stigma melanic. Hind wing yellow basally
infuscate apically. Metasoma yellowish or-
ange except the last four segments yellow-
ish orange or melanic. Ovipositor yellow-
ish orange except apical eighth melanic.

6 . — Essentially as the female.

Material examined. — Holotype. Panama:
9, Paraiso, Canal de Panama, host: Para-
thyris perspicilla Stoll (USNM). Another
specimens reviewed. Panama: 9 , Barro Col-
orado Is., 9°9'N 79°51'W, 2-9.X.1996, J.
Pickering (UK); Costa Rica: 29, 6, Prov.
Limon, Sector Cerro Cocori, Finca de E.
Rojas, 150m, 26.vi-16.vii. 1992, 12-
31.viii.1992, ii.1993, E. Rojas (INBio); 9,
Prov. Puntarenas, Est. Sirena, P.N. Corco-
vado, 0-1 00m, ii.1992, G. Rodriguez (IN-
Bio); 9, Prov. Puntarenas, Vuelta Cam-
pana, R. Terraba, 100-150m, 10-
31.viii.1992, S. Rojas (INBio); Peru: Loreto,
Pucallpa, 24. vi. 1963, J. M. Schunke
(BMHN).

Distribution. — This species is distributed
from Costa Rica and Panama in Central
America to Peru in South America.

Biology. — Larvae of Sesioctonus parathyr-
idis were reported as parasitoids in larvae
of the arctiid Parathyris perspicilla (Viereck
1914).

Sesioctonus peruviensis Briceno, sp. n.

Diagnosis. — S. peruviensis is known only
from one specimen, however, can be dis-
tinguished from all other species by the
following combination of characters: max-
illa with four palpomeres and labium with
three, occipital tubercles present, occiput
excavated, foretibia with spines. Speci-
mens of peruviensis could be confused
with S. garciai specimens, however they
are separated for the presence of epicne-
mial carinae bilobed in S. peruviensis,
which is straight in S. garciai.

Description. — 9. Length. Body, excluding
ovipositor, 5.5 mm. Head: Flagellum with
31 flagellomeres. Interantennal space with
rounded longitudinal keel. Antennal sock-

268

Journal of Hymenoptera Research

ets moderately excavated. Face without
median longitudinal carina. Genae mod-
erately expanded posteroventrally. Occip-
ital tubercles present. Occiput excavated.
Mandible concave, outer tooth longer than
inner tooth. Maxilla with 4 palpomeres.
Third and fourth labial palpomeres com-
pletely fused. Mesosoma: Subpronope tri-
angular. Longitudinal carinae of scutellar
depression absent. Scutellum convex. Me-
dian areola of metanotum smooth; with-
out median longitudinal carina; and with
lateral carinae present and not meeting
posteriorly. Propodeum convex, median
longitudinal carina absent. Epicnemial ca-
rina sharp, complete, bilobed medially
(between forecoxae). Foretibia spines pre-
sent. Midtibia with 10 spines. Hind tibia
with 18 spines. Hind femur 3.27 times as
long as wide. (RS + M)a vein of forewing
complete. 3RSa vein of forewing present.
2-1 A vein of hind wing not tubular. CUb
vein of hind wing not tubular. Hind wing
with 4 hamuli. Metasotna: Median tergite
of first metasomal segment without pair of
lateral longitudinal carinae. First metaso-
mal median tergite without depression
posterad spiracle. Ovipositor 5 mm. Color:
Head melanic. Antenna melanic. Maxil-
lary palpomeres yellowish orange except
basal two palpomeres melanic. Labial pal-
pomeres melanic. Mesosoma yellowish or-
ange. Forelegs yellowish orange except
trochantellus melanic, femur yellowish or-
ange, melanic basally and tarsus mostly
yellowish orange, but apical tarsomere
melanic. Midlegs yellowish orange except
trochanter, tibia and tarsus melanic Hind-
leg mostly melanic except coxa yellowish
orange and femur yellowish orange, me-
lanic basally. Forewing infuscate with hy-
aline spots. Stigma melanic. Hind wing
entirely infuscate. Metasoma yellowish or-
ange. Ovipositor yellowish orange except
apical eighth melanic.

6 . — Unknown.

Material examined. — Holotype. Peru: 9,
Quincemil, 750m near Marcapata, 20-
30.X.1962, Luis Pena (AEI).

Distribution. — Known only from Marca-
pata, region of Peru.

Etymology. — This species is named after
the country of the holotype specimen.

Sesioctonus qui Briceno, sp. n.

Diagnosis. — S. qui is distinguised from
all other species for the following combi-
nation of characters: occipital tubercles ab-
sent, interantennal space with a sharp lon-
gitudinal keel, gena strongly expanded
posteroventrally, median areola of meta-
notum with longitudinal rugosities and
first metasomal median tergite with de-
pression posterad spiracle.

Description. — 9. Length. Body, excluding
ovipositor, 10-15 mm (10.0). Head: Flagel-
lum with 45 flagellomeres. Interantennal
space with sharp longitudinal keel. Anten-
nal sockets deeply excavated. Face with-
out median longitudinal carina. Genae
strongly expanded posteroventrally. Oc-
cipital tubercles absent. Occiput not exca-
vated. Mandible flat, outer tooth not lon-
ger than inner tooth. Maxilla with 4 pal-
pomeres. Third and fourth labial palpo-
meres completely fused. Mesosoma:
Subpronope triangular. Longitudinal ca-
rinae of scutellar depression absent. Scu-
tellum convex. Median areola of metano-
tum with longitudinal rugosities; without
median longitudinal carina; and lacking
lateral carinae. Propodeum convex, medi-
an longitudinal carina absent. Epicnemial
carina, blunt, incomplete laterally, or com-
pletely absent, bilobed medially (between
forecoxae). Foretibial spines absent. Mid-
tibia with 5-8 spines (5). Hind tibia with
14-16 spines (14). Hind femur 4.27 times
as long as wide. (RS + M)a vein of fore-
wing complete. 3RSa vein of forewing ab-
sent. 2-1A vein of hind wing tubular. CUb
vein of hind wing not tubular. Hind wing
with 8 hamuli. Metasoma: Median tergite
of first metasomal segment without pair of
lateral longitudinal carinae. First metaso-
mal median tergite with depression pos-
terad spiracle. Length width ratio of first
metasomal median tergite 0.84. Ovipositor

Volume 12, Number 2, 2003

269

10-12 mm (10.0). Color: Head melanic.
Antenna melanic. Maxillary and labial
palpomeres yellowish orange except basal
two palpomeres melanic. Mesosoma yel-
lowish orange. Forelegs yellowish orange.
Midlegs yellowish orange. Hind leg me-
lanic except coxa yellowish orange but
melanic laterally, hind femur yellowish
orange, hind tibia yellowish orange in bas-
al half, melanic apically. Forewing banded
from base, yellow, infuscate, yellow, in-
fuscate. Stigma yellowish orange. Hind
wing yellow basally infuscate apically.
Metasoma yellowish orange with the last
four tergites melanic. Ovipositor yellow-
ish orange.

6. — Male exhibits a color darker than
females, showing the body brownish.
However, only one male of this specie was
examined for this revision and another fu-
ture observations are necessary.

Material examined. — Holotype: Venezuela:
9, Aragua, El Limon, 450m, 25.vi.1978, luz
de mercurio, Francisco Fernandez Yepez
col (MIZA). Paratypes. Brasil: 2$, Sao Pau-
lo, Teodoro Sampaio, xii.1977, F. M. Oliv-
eira (CNC) (BMNH); Peru: 9, 6, Loreto,
Pucallpa, 19.iv.1962, vi.1965, J. M. Schunke
(BMNH).

Distribution. — This species is distributed
from Venezuela until Brazil and Peru in
South America. Its presence in Colombia,
Ecuador and Bolivia is probable.

Etymology. — The species name qui is an
arbitrary combination of letters.

Sesioctonns theskelos Briceno, sp. n.

Diagnosis. — S. theskelos can be distin-
guished for the following characters com-
bination: lacking of occipital tubercles,
maxilla with 5 palpomeres and labial with
4, median longitudinal carina of propo-
deum present, and median tergite of first
metasomal segment with pair of lateral
longitudinal carinae.

Description. — 9. Length. Body, excluding
ovipositor, 7 mm. Head: Flagellum with 35
flagellomeres. Interantennal space with
rounded longitudinal keel. Antennal sock-

ets moderately excavated. Face without
median longitudinal carina. Genae mod-
erately expanded posteroventrally. Occip-
ital tubercles absent. Occiput not excavat-
ed. Mandible concave, outer tooth of man-
dible not longer than inner tooth. Maxilla
with 5 palpomeres. Third and fourth labial
palpomeres not fused. Mesosoma: Sub-
pronope triangular. Longitudinal carinae
of scutellar depression absent. Scutellum
convex. Median areola of metanotum
smooth; without median longitudinal ca-
rina; and with lateral carinae present and
not meeting posteriorly. Propodeum con-
vex, median longitudinal carina present.
Epicnemial carina sharp, complete later-
ally, bilobed medially (between forecox-
ae). Foretibia spines absent. Midtibia with
6-9 spines (9). Hind tibia with 12-15
spines (12). Hind femur 3.5 times as long
as wide. (RS+M)a vein of forewing com-
plete. 3RSa vein of forewing present or ab-
sent. 2-1A vein of hind wing tubular or
not tubular. CUb vein of hind wing not
tubular. Hind wing with 3-A (4) hamuli.
Metasoma: Median tergite of first meta-
somal segment with pair of lateral longi-
tudinal carinae. First metasomal median
tergite without depression posterad spi-
racle. Length width ratio of first metaso-
mal median tergite 0.94-1.12 (0.94). Ovi-
positor 5-6 mm (5.5). Color: Head melanic
sometimes with a spot yellowish orange
on the front. Antenna melanic. Maxillary
and labial palpomeres yellowish orange.
Mesosoma yellowish orange. Forelegs yel-
lowish orange. Midlegs yellowish orange
except tarsus melanic. Hindleg melanic
except coxa yellowish orange, or yellow-
ish orange but melanic laterally; and fe-
mur yellowish orange, sometimes melanic
distally. Forewing entirely infuscate. Stig-
ma melanic. Hind wing entirely infuscate.
Metasoma entirely yellowish orange or
yellowish orange with the last four ter-
gites melanic posteriorly. Ovipositor yel-
lowish orange.

6 . — Unknown.

Material examined. — Holotype. Brasil: 9,

270

Journal of Hymenoptera Research

Matogrosso, Sinop, x.1975, Trampa mal-
aise, M. Alvarenga (CNC). Paratypes.
Brasil: 9, Matogrosso, Sinop, x.1975,
Trampa malaise, M. Alvarenga (CNC); Ec-
uador: 9, Coca, v.1992, Luis Pena (AEI).

Distribution. — This species is known
only from Brazil and Ecuador.

Etymology. — The species name tJieskelos
means wonderful.

Sesioctomis venezuelensis Briceno, sp. n.

Diagnosis. — S. venezuelensis is the only
one species that shows five longitudinal
carinae on the scutellar depression, in
combination with the presence of occipital
tubercles and occiput excavate.

Description. — 8. Length. Body, excluding
ovipositor, 8 mm. Head: Flagellum with
broken after flagellomere 28. Interantennal
space with rounded longitudinal keel. An-
tennal sockets moderately excavated. Face
without median longitudinal carina. Ge-
nae not expanded posteroventrally. Occip-
ital tubercles present. Occiput excavated.
Mandible concave. Maxilla with 4 pal-
pomeres. Third and fourth labial palpo-
meres completely fused. Mesosoma: Sub-
pronope oval. Longitudinal carinae of scu-
tellar depression present. Scutellum flat.
Median areola of metanotum smooth;
without median longitudinal carina; and
with lateral carinae present and not meet-
ing posteriorly. Propodeum flat, median
longitudinal carina absent. Epicnemial ca-
rina sharp, complete, bilobed medially
(between forecoxae). Foretibia spines ab-
sent. Midtibia with 7 spines. Hind tibia
with 16 spines. Hind femur 3.5 times as
long as wide. (RS+M)a vein of forewing
complete. 3RSa vein of forewing present.
2-1 A vein of hind wing not tubular. CUb
vein of hind wing not tubular. Hind wing
with 5 hamuli. Metasoma: Median tergite
of first metasomal segment without pair of
lateral longitudinal carinae. First metaso-
mal median tergite without depression
posterad spiracle. Length width ratio of
first metasomal median tergite 1.28. Color:
Head melanic. Antenna melanic. Maxil-

lary and labial palpomeres yellowish or-
ange. Mesosoma melanic. Forelegs yellow-
ish orange except coxa melanic. Midlegs
yellowish orange except coxa, trochanter
and trocantellus melanic. Hindleg melanic
except femur melanic but yellowish or-
ange apically. Forewing banded from
base, yellow, infuscate, yellow, infuscate.
Stigma melanic. Hind wing banded from
base, yellow, infuscate, yellow, infuscate.
Metasoma yellowish orange with third
tergum yellowish orange but median ter-
gite melanic in posterior quarter and
fourth and fifth tergum melanic.

9 .—Unknown.

Material examined. — Holotype. Venezuela:
8 , Aragua Est. Exp. Cataurito, 32km E Vil-
la de Cura, 1100m, 28.vi.1983, O. S. Flint,
Jr. (CNC).

Distribution. — This species is known
only from the Aragua region of Venezue-
la.

Etymology. — This species is named after
the country of the holotype specimen.

ACKNOWLEDGMENTS

I am sincerely grateful to the entomological collec-
tions that lent the specimens for this research. Special
acknowledgment to Dr. Michael Sharkey for his im-
portant advising and to Ing. Quintin Arias for assis-
tance with graphics software. Jose Clavijo, Jose Luis
Garcia and Jurg De Marmels did important correc-
tions for the first manuscript. The "Centro de Mi-
croscopia Electronica de la Facultad de Agronomia-
UCV" (CENMEFA- Venezuela) prepared specimens
for SEM and took the photomicrografhs. Support for
this investigation was provided by FONACIT (Project
SI-2000000479), by the Consejo de Desarrollo Cientf-
fico, Humanistico y Tecnologico (CDCHT-028-AG-
2001) of the Universidad Centroccidental "Lisandro
Alvarado" (Barquisimeto- Venezuela) and by the Fun-
dacion Cientifica y Tecnologica of the Aragua state
(FUNDACITE- Aragua- Proyecto 2000-FHR-02-05-02-
1). Specimens from Colombia were provided by
M.Sharkey through with the support of NSF grant
DEB9972024.

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Sharkey, M. J. 1988. A taxonomic revision of Alaba-
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Sharkey, M. J. 1992. Cladistic and tribal classification
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Sharkey, M. J. 1997. Subfamily Agathidinae. In Whar-
ton, R. A.; P. M. Marsh and M. J. Sharkey. Manual
of the New World genera of the family Braconidae
(Hymenoptera). Special publication of the Inter-

national Society of Hymenopterists. Number 1.
Washington D. C. 439 p.

Viereck, H. L. 1912. New genus and species of Hy-
menoptera of the family Braconidae from Pana-
ma. Smithsonian Miscellaneous Collections. 59: 1-2.

Viereck, H. L. 1914. Type species of Ichneumon-flies.
Bulletin of the United States National Museum. 83:
1-186.

Wharton, R. A., P. M. Marsh, and M. J. Sharkey. 1997.
Manual of the Neiv World genera of the family Bra-
conidae (Hymenoptera). Special Publication of the
International Society of Hymenopterists. Num-
ber 1. Washington, D. C. 439 p.

J. HYM. RES.

Vol. 12(2), 2003, pp. 272-277

The Effect of Body Size on Male-Male Combat in the Parasitoid
Wasp Melittobia digitata Dahms (Hymenoptera: Eulophidae)

Christopher S. Hartley and Robert W. Matthews

University of Georgia, Department of Entomology, Athens, GA 30602-2603, USA,
CSH email: chartley@uga.edu; RWM email: rmatthew@uga.edu

Abstract.— The parasitic wasp Melittobia digitata Dahms (Hymenoptera: Eulophidae) is a gregar-
ious external parasitoid of various insects, primarily solitary wasps and bees. Males of M. digitata
commonly engage in fierce, often fatal, fights with other males. The mandibles are the main
weapons used, and injuries inflicted vary greatly in severity from loss of appendages to death.
We investigated the effect of size differences between winning, losing, and non-fighting males
and whether body size was related to being a fighter or non-fighter. The head width and tibia
length of fighting and non-fighting males were measured. Winning males (21 of 29 pairs) were
found to be larger than losing males, and fighting males (winners and losers combined) were
found to be larger than non-fighting males (8 pairs). Mandible lengths of a subset of all males (15
fighters, 12 non-fighters) were subsequently measured; only fighter's left mandible length was
significantly correlated with head width. The possibility that two behavioral forms (fighters and
non-fighters) exist is considered but will require further experiments to resolve.

Ritualized fighting over food, territory, the swollen inflorescences of various fig

mates and other resources occurs between species, and fights occur between well-ar-

males in the majority of animal species, mored flightless males that pursue newly

Many studies have emphasized the re- emerged females. The majority of males

strained nature of male fights by showing do not emerge from the fig in which they

that the majority of fights end peacefully are born though emergence is more com-

with neither combatant being injured, mon in some species (Bean and Cook

even in species where the males possess 2001). Males have also been shown to

large and dangerous weapons (Maynard have a strong attraction to the fig in which

Smith and Price 1973; Maynard Smith they were reared (Frank 1985), which

1982). In addition, fighting males often makes dispersal unlikely. Since potential

display striking dimorphisms such as mating opportunities are limited, fights

those found in certain beetles (Forsyth and between males are fierce and result in

Alcock 1990; Zeh et al. 1992; Goldsmith many fatalities.

and Alcock 1993) and mites (Saito 1990, Melittobia digitata Dahms (Hymenop-

1995). Instances of fatal fighting have been tera: Eulophidae) is a gregarious external

observed in some animal species. Where parasitoid of many different insects, but

they do occur, fatal fights usually involve its principal hosts are solitary wasps and

opportunities to mate, and they are limit- bees. In the southeastern United States, M.

ed to species where males have limited digitata is most commonly found attacking

opportunities to mate (Enquist and Leimar the mud dauber wasp, Trypoxylon politum

1990). Fatal fighting has been well docu- Say (Hymenoptera: Sphecidae). A female

mented in many fig wasps (Hamilton Melittobia enters a Trypoxylon cocoon be-

1979; Murray 1987; Bean and Cook 2001). fore it is sealed and waits until the host

These wasps spend their entire life within transforms into a prepupa before ovipos-

Volume 12, Number 2, 2003 273

iting hundreds of eggs directly onto the culturing in the laboratory does not alter
prepupa's cuticle (Dahms 1984; Gonzalez Melittobia behavior (Assem and Jachmann
and Teran 2001). Upon hatching, the gre- 1999), so intense fighting is not likely to
gariously developing Melittobia larvae be an artifact of mass rearing,
consume the host, complete their devel- The occurrence of fatal fighting in both
opment and begin to emerge in approxi- fig wasps and Melittobia is unexpected be-
mately 20 days, depending on tempera- cause males are normally fighting their
ture. The average brood size for M. digitata brothers. Hamilton (1979) suggested that
resulting from one female foundress is 522 fighting behavior would not exist where a
young. Males and females emerge togeth- male's rival has a high chance of being a
er, but the sex ratio is extremely female brother. Recent work with fig wasps, how-
biased — about 98% female (J.M. Gonzalez, ever, found no relationship between relat-
personal communication). Females char- edness of males and fighting behavior. It
acteristically mate once, usually soon after was found instead that the level of fatal
emergence, and one male may mate with fighting was negatively correlated with fu-
numerous females in his lifetime. Mated ture mating opportunities (West et al.
females then chew their way out of the 2001). A similar situation exists in Melit-
host's cell and disperse to search for new tobia where males have little chance of fu-
hosts (Dahms 1984). ture mating opportunities since they are

Sexual dimorphism is extreme in Melit- not likely to disperse.
tobia. Adult males possess vestigial eyes, The objective of this study was to de-
short non-functional wings, enlarged an- termine if size differences exist in M. dig-
tennal scapes, and mandibles with well- itata between winners and losers of fights
developed teeth. The blind males wander and between fighting and non-fighting
freely inside the host's cocoon until they males. We hypothesized that winners
encounter a female or another male. En- would be larger than losers and that fight-
counters with females instigate courting ers would be larger than non-fighters,
and mating behaviors, and virgin females
often gather in groups around males to MATERIALS AND METHODS

await mating (Gonzalez et al. 1985; Con- Melittobia digitata cultures were reared

soli et al. 2002). Encounters with other in an incubator at 25°C on T. politum pre-

males quickly escalate into a grappling pupae in small plastic boxes (5cm X 2.5cm

contest where the males interlock their x 1.8cm) with tightly fitting lids. Males

legs and struggle briefly with each other, were removed from cultures as pupae and

Following these bouts, the males will ei- isolated in Carolina^ clear Deep Well Pro-

ther separate or attempt to use their man- jection Slides (25 mm diameter, 2 mm

dibles to tear at the body of the opponent, deep). This isolation ensured that a male's

These fights often lead to loss of append- age and prior mating and fighting expe-

ages and death in one or both fighters rience could be controlled. No data were

(Dahms 1984). Inside naturally parasitized recorded on male emergence time relative

mud dauber cocoons, one routinely finds to other males from a particular culture

the remains of several males, many dis- nor from which culture a given male

membered. In most laboratory cultures of emerged. Thus, males used in the experi-

M. digitata, males grapple and fight with ments can be regarded as arbitrarily se-

little provocation, and these contests fre- lected from among a range of males avail-

quently end with the death of one or both able.

combatants. In other cultures, we have When the males isolated in the depres-

found many males alive with no injuries sion slides emerged, the date of their

and no evidence of fighting. Long term emergence was recorded. Eighty-seven

274

Journal of Hymenoptera Research

Table 1. Morphometric measurements of M. digitata males.

Total number

Winners
Losers
Fighters
Non-Fighters

21
21
42
16

* Values in mm ± Standard Deviation.

Mean tibia length*

0.25 ± 0.03
0.23 ± 0.03
0.24 ± 0.03
0.20 ± 0.03

Mean head width*

0.35 ± 0.04

0.34 ± 0.04

0.35 ± 0.04

0.29 ± 0.03

males were kept isolated and observed
daily, and their date of death was record-
ed. These males served as controls for the
following experiment.

Twenty-nine newly emerged male pairs
were formed in the depression slides by
opening the two individual slides and us-
ing a paintbrush to move one of the males
into the other male's slide. Because of the
difficulty of marking individuals, it was
not possible to track which male was res-
ident versus intruder in the pairings. Once
each pair had been formed, the slides
were not opened again until after both
males had died. Each pair was observed
daily, and the date of each male's death
was recorded.

After death, each male was examined,
and any obvious injury (e.g., loss of ap-
pendages, body wounds) was recorded.
The head and right front tibia of each male
were then mounted temporarily in glyc-
erol on standard glass microscope slides
and measured under 50 X magnification
using an ocular micrometer. Both males of
one pair were mounted on the same slide
so that winners and losers could be asso-
ciated. To avoid crushing the specimens,
pieces of 61b. test nylon fishing line were
placed around them to elevate the cover
slip.

To test whether head width is correlated
with mandible length, 15 fighter heads
and 12 non-fighter heads were arbitrarily
selected, and their mandibles were dis-
sected. Both left and right mandibles were
mounted on microscope slides and their
maximum length was measured.

Sign tests were used to analyze winner

versus loser data so that both males of one
fighting pair could be compared against
each other. Mann-Whitney U tests were
used to analyze fighter versus non-fighter
data. Spearman R Correlations were used
for all correlations. A P value of 0.05 was
taken as the critical value for establishing
significance. Analyses were done using
STATISTICA 6 © StatSoft, Inc.

RESULTS

In 21 of the pairs, one male killed the
other in a fight. These males were termed
fighters. In eight of the pairs, the males
were never observed to come into contact
with each other, and after death neither
male was found to have lost appendages
or incurred wounds to the body. We con-
cluded in these cases that no fighting had
occurred, and these males were recorded
as non-fighters.

Table 1 shows the measurement data for
all males.

Winners had significantly longer tibia
(Z = 2.29; P = 0.022), but head widths of
winners and losers did not differ signifi-
cantly (Z = 1.21; P = 0.228). The tibia
length of all fighters (winners and losers
combined) was significantly longer than
the non-fighters' tibia length (U = 134.0;
Z - 3.58; P < 0.001), and fighters' heads
were significantly wider than non-fight-
ers' (U - 88.5; Z = 4.34; P < 0.001). Head
width and tibia length for all males (fight-
er and non-fighter) were significantly cor-
related (p = 0.665; P < 0.001).

The only significant correlation between
head width and mandible length was
found for fighters' left mandibles (p =

Volume 12, Number 2, 2003

275

14

12

10

3

£ 6

■ Fighters

■ Non-fighters

1

«- CM

o d

CO

o

d

d

CD

d

d

CM t-
^ CM

cm co

CM CM

odd

CM

CM

CD
CM

CM

co

CM

Tibia Length (mm)

Fig. 1. Frequency distribution for tibia length comparing all fighters and non-fighters (n = 58).

0.692; P = 0.004). Fighters' right mandi-
bles were not significantly correlated with
head width (p = 0.351; P = 0.200). Non-
fighters' left mandibles were not signifi-
cantly correlated with head width (p =
0.507; P = 0.092), and non-fighters' right
mandibles were not significantly correlat-
ed with head width (p = 0.452; P = 0.140).
Differences in size between fighters and
non-fighters and the obvious behavioral
differences suggested the possibility that
the Melittobia male population could be di-
morphic. To explore this possibility, fre-
quency distributions of the measurements
of tibia length and head width were pre-
pared (Figs. 1 and 2). If a dimorphism ex-
ists, a bimodal curve is expected. The
graph of tibia length frequency reveals
only one peak for both fighters and non-
fighters, and this peak occurs at 0.23mm.
The graph of head width frequencies sug-
gests the presence of two peaks — one for
non-fighters at 0.28mm and one for fight-
ers at 0.37mm. There is, however, consid-

erable overlap, and values for fighters and
non-fighters occur at both ends of the
scale.

DISCUSSION

We predicted that winners would be the
larger males. The results revealed that
winners were larger than losers based on
their tibia length measurements, although
their head widths did not differ. The cor-
relation between head width and tibia
length suggests that relative size of either
is likely to be a good predictor of overall
body size, which in turn is related to fight-
ing success. Mandible size has been often
discussed in relation to fighting fig wasps
(Bean and Cook 2001). The frequent asym-
metry between right and left mandible
lengths in our study was unexpected, and
may be interesting to pursue.

The existence of non-fighter males, in
which paired males never engaged in a
fight even though they were isolated to-
gether for their whole life, raises the pos-

276

Journal of Hymenoptera Research

Fig. 2. Frequency distribution for head width comparing all fighters and non-fighters (n = 58).

sibility that males exist in two behavioral
morphs. Freeman and Ittyeipe (1982) de-
scribed two morphologically distinct male
morphs in Melittobia hawaiiensis Perkins ( =
M. australica Girault): a larger morph with
ocelli and a smaller morph without ocelli.
However, we found that all of our M. dig-
itata males, large and small, had fully pig-
mented ocelli. The frequency graphs of
head width and tibia length (Figs. 1 and
2), while showing a trend towards a bi-
modal distribution for each trait, also re-
veal that a wide range of sizes exists in
both fighting and non-fighting males.
Overall, most of the larger males became
fighters while most of the smaller males
never engaged in fights, but there were
obvious exceptions. Perhaps fighting and
non-fighting are conditional rather than
fixed traits. Alternatively, there could be
culture or lineage specific effects on the
likelihood of a male becoming a fighter or
non-fighter. These questions will require
further investigation.

Abe et al. (2003) found that when an
emerged male and a pupal male were
placed together, the already emerged male
usually killed the pupal male at or im-
mediately after eclosion, but they did not
record sizes of any of the males in their
experiment. We have noted that the first
males to emerge are generally larger most
likely due to better food quality and quan-
tity, and small males emerge later in the
culture's life, when the host is covered
with developing pupae (unpublished
data). Males of different fig wasp species
are known to exist in a wide variety of
body shapes, each with a different fighting
propensity, and each is adapted to court-
ing females in a different way (Murray
1990). If fighting behavior in Melittobia is
linked to size, then perhaps a similar sit-
uation occurs with small, non-fighting
males being better adapted to maneuver-
ing amongst the developing pupae in the
tightly packed confines of a Trypoxylon co-
coon. They could avoid the stress of fight-

Volume 12, Number 2, 2003

277

ing by staying hidden, but they could still
court females. Repeating these experi-
ments using males from one culture and
tracking relative emergence times could
help to determine if males that emerge
early tend to be larger and become fight-
ers and males that emerge late tend to be
smaller and become non-fighters.

ACKNOWLEDGMENTS

Jorge M. Gonzalez and Leif Deyrup provided in-
valuable comments on the experimental design and
the manuscript. We also thank Jan Matthews, David
Jenkins, Christian Torres, and LuAnn Brown for all
of their help and advice. This study was conducted
as a senior Honor's Thesis at the University of Geor-
gia, and was supported in part by NSF Grant
0088021, R. W. Matthews, Principal Investigator.

LITERATURE CITED

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2003. Extremely female-biased sex ratio and le-
thal male-male combat in a parasitoid wasp, Mel-
ittobia australica (Eulophidae). Behavioral Ecology
14: 6-11.

Assem, J. van den, and F. Jachmann. 1999. Changes
in male perseverance in courtship and female
readiness to mate in a strain of the parasitic wasp
Nasonia vitripennis over a period of 20+ years.
Netherlands Journal of Zoology 49: 125-137.

Bean, D., and J. M. Cook. 2001. Male mating tactics
and lethal combat in the nonpollinating fig wasp
Sycoscapter australis. Animal Behaviour 62: 535-
542.

Consoli, F. L., H. J. Williams, S. B. Vinson, R. W. Mat-
thews, and M. F. Cooperband. 2002. trans-Ber-
gamotenes — Male pheromone of the ectoparasi-
toid Melittobia digitata. journal of Chemical Ecology
28: 1675-1689.

Dahms, E. C. 1984. A review of the biology of species
in the genus Melittobia (Hymenoptera: Eulophi-
dae) with interpretations and additions using ob-
servations on Melittobia australica. Memoirs of the
Queensland Museum 21: 337-360.

Enquist, M., and O. Leimar. 1990. The evolution of
fatal fighting. Animal Behaviour 39: 1-9.

Forsyth, A., and J. Alcock. 1990. Female mimicry and
resource defense polygyny by males of a tropical

rove beetle Leistotrophus versicolor Coleoptera Sta-
phylinidae. Behavioral Ecology and Sociobiology 26:
325-330.

Frank, S. A. 1985. Are mating and mate competition
by the fig wasp Pegoscapus assuetus (Agaonidae)
random within a fig? Biotropica 17: 170-172.

Freeman, B. E., and K. Ittyeipe. 1982. Morph deter-
mination in Melittobia, a eulophid wasp. Ecologi-
cal Entomology 7: 355-363.

Goldsmith, S. K., and J. Alcock. 1993. The mating
chances of small males of the cerambycid Trach-
yderes mandibularis differ in different environ-
ments (Coleoptera: Cerambycidae). journal of In-
sect Behavior 6: 351-360.

Gonzalez, J. M., and J. B. Teran. 2001. Dispersion,
busqueda y acceso al hospedador por Melittobia
acasta (Hymenoptera: Eulophidae). Boletin del
Centro de Investigations Biologicas 35: 52-64.

Gonzalez, J. M., R. W. Matthews, and J. R. Matthews.
1985. A sex pheromone in males of Melittobia aus-
tralica and Melittobia femorata (Hymenoptera: Eu-
lophidae). Florida Entomologist 68: 279-286.

Hamilton, W. D. 1979. Wingless and fighting males
in fig wasps and other insects, pp. 167-220. In
M.S. Blum and N.A. Blum (eds.), Sexual Selection
and Reproductive Competition in Insects. Academic
Press, New York.

Maynard Smith, J. 1982. Evolution and the Theory of
Games. Cambridge University Press, Cambridge.

Maynard Smith, J., and G. R. Price. 1973. The logic of
animal conflict. Nature 246: 15-18.

Murray, M. G. 1987. The closed environment of the
fig receptacle and its influence on male conflict
in the Old World fig wasp, Philotrypesis pilosa.
Animal Behaviour 35: 488-506.

Murray, M. G. 1990. Comparative morphology and
mate competition of flightless male fig wasps.
Animal Behaviour 39: 434-443.

Saito, Y. 1990. Factors determining harem ownership
in a subsocial spider-mite (Acari, Tetranychidae).
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Saito, Y. 1995. Clinal variation in male-to-male antag-
onism and weaponry in a subsocial mite. Evolu-
tion 49: 413-417.

West, S. A., M. G. Murray, C. A. Machado, A. S. Grif-
fin, and E. A. Herre. 2001. Testing Hamilton's
rule with competition between relatives. Nature
409: 510-512.

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86-96.

J. HYM. RES.
Vol. 12(2), 2003, pp. 278-302

Morphological Variation in Opius Wesmael
(Hymenoptera: Braconidae) with an Emphasis on Nearctic Species

in the Subgenus Gastrosema Fischer

Robert R. Kula

Department of Entomology, Texas A&M University, College Station, TX, USA 77843,

email: rkula@oznet.ksu.edu

Abstract— The variability of morphological characters and their potential phylogenetic utility in
Opiinae are evaluated using Nearctic species in the subgenus Gastrosema Fischer. Diagnoses for
three species groups within Nearctic Gastrosema are provided, and 43 Nearctic species in Opius
Wesmael are placed in the species groups. Two new Nearctic species in Gastrosema are described:
Opius abbyae Kula and O. lacopitaensis Kula. Five previously described Nearctic species in
Gastrosema are redescribed: O. castaneigaster Fischer, O. intermissus Fischer, O. oscinidis (Ashmead),
O. salmonensis Fischer, and O. striativentris Gahan. Opius gracillariae Gahan is designated a junior
synonym of O. striativentris.

Opiinae is a very large subfamily in Bra- ical limits of these subgenera need to be

conidae with approximately 1,500 de- evaluated. This research begins to address

scribed species worldwide (Wharton these problems by performing a detailed

1997b). All valid rearing data suggest that assessment of intraspecific morphological

species in Opiinae are strict koinobiont en- variation for Nearctic species in Gastrose-

doparasitoids of cyclorrhaphous Diptera ma.

(Wharton 1999). Opius Wesmael is the Other than Fischer's (1977) revision of
largest genus in Opiinae, and over 1,000 New World Opiinae, there has been little
species are currently placed in Opius sensu work on Nearctic species in Gastrosema.
Fischer (1972). In a series of regional re- Gahan (1915) described two Nearctic spe-
visions, Fischer (1972, 1977, 1987) segre- cies currently included in Gastrosema and
gated species in Opius into 27, 30, and 34 provided a key to the North American
subgenera, respectively. Most of the sub- species in Oprius. Fischer (1964, 1970) de-
genera are broadly defined and lack fea- scribed several Nearctic species in Opius
tures that clearly characterize them as that were subsequently placed in Gastro-
monophyletic. Further, the defining fea- sema by Fischer (1977). Marsh (1974) syn-
ture of several subgenera, including Gas- onymized two Nearctic species in Gastro-
trosema Fischer, Merotrachys Fischer, Phae- sema, but insufficient material was exam-
drotoma Forster sensu Fischer (1972), and ined to assess intraspecific morphological
Tolbia Cameron sensu Fischer (1972), is the variation. Additional specimens must be
presence or absence of sculpture associat- acquired and compared with the holo-
ed with specific anatomical features. Intra- types to confirm Marsh's synonymies. Van
specific variation of sculpture has never Achterberg (1997) transferred O. pumilio
been assessed for species in these subgen- Wesmael (the type species for Gastrosema)
era, and preliminary sorting indicates that to Phaedrotoma sensu van Achterberg and
certain species fit the concept of more than Salvo (1997). This action suggests that van
one subgenus. Therefore, the morpholog- Achterberg (1997) treated Gastrosema Fi-

Volume 12, Number 2, 2003

279

scher 1972 as a junior synonym of Phae-
drotoma Forster 1862, although this was
not explicitly stated. Since only some spe-
cies currently classified as Gastrosema pos-
sess the characters used to define Phaedro-
toma sensu van Achterberg and Salvo
(1997), synonymy seems inappropriate.
Additionally, Phaedrotoma sensu van Ach-
terberg and Salvo (1997) is partially de-
fined by the absence of a basal mandibular
tooth which the authors regarded as ple-
siomorphic. It appears that Phaedrotoma
sensu van Achterberg and Salvo (1997) has
not been defined as a monophyletic
group, and I see no advantage in accept-
ing the changes proposed by van Achter-
berg (1997) over Fischer's (1972) concept
of Gastrosema.

MATERIALS AND METHODS

Specimens used in this study were bor-
rowed from several North American mu-
seums and collections. The American En-
tomological Institute (AEI), the California
Academy of Sciences (CAS), the Canadian
National Collection of Insects (CNCI), the
Insect Research Collection at the Univer-
sity of Wisconsin-Madison (IRCW), the
Museum of Comparative Zoology at Har-
vard University (MCZ), the insect collec-
tion at Texas A&M University (TAMU),
and the United States National Museum
of Natural History (USNM) provided de-
termined and undetermined opiines. In
addition to the borrowed material, two
opiine species treated in this study were
reared from isolated puparia. Dr. Sonja J.
Scheffer provided several hundred speci-
mens of O. striativentris Gahan reared
from five species of Phytomyza Fallen min-
ing the leaves of seven species of Ilex L.
(hollies). Eighteen specimens of an unde-
scribed opiine were reared by the author
from a species of Calycomyza Hendel min-
ing the leaves of Helianthus annuus L. (sun-
flower).

Species treated in this study can be
identified to Opiinae using Sharkey (1997)
and to Opius using Wharton (1997a). Sev-

eral hundred Nearctic species in at least
11 subgenera have not yet been examined
but potentially fall into the species groups
treated in this paper. As a result a com-
prehensive key to the Nearctic species in
each species group cannot be provided at
this time. Therefore, a detailed diagnosis
is provided for each species group and
species to facilitate their identification.

Descriptive terminology for anatomical
features, including wing cells and veins, is
based on Sharkey and Wharton (1997).
Terminology used to describe surface
sculpture follows Harris (1979) and Shar-
key and Wharton (1997). Two anatomical
terms used throughout this work are not
found in Harris (1979) or Sharkey and
Wharton (1997). Both terms must be de-
fined and illustrated to provide maximum
clarification. The anterior pronotal furrow
is a groove located at the antero-lateral
edge of the pronotum just posterior to the
propleural flange, and the posterior me-
sopleural furrow is a groove located at the
posterior edge of the mesopleuron just an-
terior to the metapleuron (Fig. 1).

Several measurements were taken to
quantitatively characterize each species
described or redescribed in this study
(Figs. 2-7). Two criteria were used to se-
lect measurements: 1) measurements his-
torically used in opiine species descrip-
tions and 2) measurements that describe
the general size and shape of each species.

MORPHOLOGICAL VARIATION

Results from the analysis of intraspecific
morphological variation provide a basis
for discussing character variability and
their potential phylogenetic utility in Opi-
inae. The following section is an assess-
ment of characters potentially useful for
supporting monophyletic groups, as well
as characters useful for defining the limits
of species.

Size and shape ofclypeus. — Fischer (1972)
used the exposure of the lab rum in frontal
view (a result of clypeus length) to par-
tially define certain subgenera. For exam-

280

Figs. 1-7. 1, Anterior pronotal furrow (arrow a) and
posterior mesopleural furrow (arrow b). 2, Head
width (HW). 3, Mesosoma width (MW) and tergite 1
width (tlW). 4, Head length (HL), eye length (EL),
eye width (EW), mesosoma length (ML), and meso-
soma depth (MD). 5, Tergite 1 length (tlL) and ex-
posed ovipositor length (EOL). 6, Hind tibia length
(HTL). 7, 2RS length (2RSL), 3RSa length (3RSaL),
and 2M length (2ML). Figs. 1 and 3-5 modified from
Fischer (1972).

pie, species that Fischer included in Sto-
mosema Fischer possess a broad clypeus
completely concealing the labrum (Fig. 8).
Alternatively, species that Fischer includ-
ed in Gastrosema have a reduced clypeus

Journal of Hymenoptera Research

exposing the labrum in frontal view (Fig.

9).

Three basic clypeal morphologies are

observed among Nearctic species in Gas-
trosema. Both O. oscinidis (Ashmead) and
O. walleyi Fischer possess a flattened or
convex clypeus with the ventral margin
lobed mesally (Fig. 10). This condition is
also commonly observed for Nearctic spe-
cies in RJwgadopsis Brethes sensu Wharton
(1987) and Thoracosema Fischer. Opius
striativentris possesses a convex, hemi-
spherical to narrowly hemielliptical
shaped clypeus with a straight to slightly
arched ventral margin (Fig. 9). This is the
most commonly observed clypeal shape
for Nearctic species in Gastrosema and is
also commonly observed for Nearctic spe-
cies in Phaedrotoma sensu Fischer (1972)
and Tolbia sensu Fischer (1972). Both O.
flaviceps Gahan and O. sahuonensis Fischer
possess a flattened, broadly hemielliptical
shaped clypeus with a straight to slightly
arched ventral margin (Fig. 11). This clyp-
eal shape is also commonly observed for
Nearctic species in Pleurosema Fischer. The
consistency of clypeus size and shape
within a species, coupled with the vari-
ability of clypeus size and shape among
groups of species, suggests that this char-
acter is potentially phylogenetically infor-
mative and merits consideration in future
phylogenetic analyses.

Shape of mandibles. — Mandible shape is
another feature Fischer (1972) used to par-
tially define certain subgenera. For exam-
ple, Opiognathus Fischer and Opiostomus
Fischer contain species with mandibles
that are abruptly widened distally to prox-
imally (basal tooth present) (Fig. 12). In
more recent revisionary work, van Ach-
terberg and Salvo (1997) used mandible
shape to split Opius sensu Into into two
genera: Opius and Phaedrotoma. Opius sen-
su van Achterberg and Salvo (1997) in-
cludes species that possess a distinct basal
tooth, while Phnedrotomn sensu van Ach-
terberg and Salvo (1997) includes species
that lack a distinct basal tooth (Fig. 13).

Volume 12, Number 2, 2003

281

VR9V

^^^^r

fee - %^-' """

m '■■

s^fuwSHi3^

■

M

WfflH

1

Ifiu^?i

..

Figs. 8-11. Size and shape of clypeus. 8, Broad and completely concealing labrum. 9, Reduced (arrow a) and
exposing labrum (arrow b). 10, Ventral margin lobed mesally. 11, Broadly hemielliptical with slightly arched
ventral margin.

Van Achterberg and Salvo (1997) explicit-
ly regarded the presence of a basal tooth
as apomorphic. Thus, if the presence of a
basal tooth is apomorphic, Phaedrotoma
sensu van Achterberg and Salvo (1997) is
partially defined by a symplesiomorphy.
The nomenclatural changes in van Achter-
berg and Salvo (1997) collapse the subge-
neric classification established by Fischer
(1972, 1977, 1987). The species currently
included in Opius sensu lato are segregated
into the putatively monophyletic genus
Opius sensu stricto or the seemingly para-
phyletic genus Phaedrotoma sensu lato. Spe-
cies with mandibles that are gradually
widened distally to proximally (Fig. 14)
cannot be unequivocally placed in either
genus. Additionally, species that exhibit
distinct similarities in clypeal shape, body

sculpture, and wing venation (e.g. O. rc-
lativus Fischer and O. salmonensis) fall into
different genera. Because of these short-
comings I see no reason to adopt the clas-
sification proposed by van Achterberg and
Salvo (1997) over Fischer (1972, 1977,
1987).

The Nearctic species of Opius sensu lato
that fall within Gastrosema vary extensive-
ly in mandibular shape. In O. lacopitaensis
Kula, new species (described below) the
basal tooth is obviously absent in most in-
dividuals but gradually widened distally
to proximally in some individuals. The ab-
sence of the basal tooth is the most com-
monly observed condition for Nearctic
species in Gastrosema. The continuous na-
ture of this character makes it verv diffi-

J

cult to code into discrete character states

-

Journal of Hymenoptera Research

Figs. 12-14. Mandibles. 12, Basal tooth distinctly present. 13, Basal tooth completely absent. 14, Gradually
widened distally to proximally.

for certain species. However, species such
as O. salmonensis consistently lack a basal
tooth, and species such as O. relativus con-
sistently possess a distinct basal tooth. Op-
ius tangens Fischer is another Nearctic spe-
cies in Gastrosema that possesses a distinct
basal tooth. Unfortunately, only one indi-
vidual was available for examination, so
intraspecific variation of mandibular
shape is not known for O. tangens. Nearc-
tic species in several subgenera exhibit
variation in mandibular shape similar to
Nearctic species in Gastrosema. PJiaedroto-
ma sensu Fischer (1972) and Pleurosema
contain species that either obviously lack
or distinctly possess a basal tooth, as well
as species with mandibular shapes that
cannot be reliably determined as basal
tooth present or absent. The results from
this study suggest that mandible shape is

useful for distinguishing between certain
species but may not be possible to code
unambiguously for phylogenetic analyses.

Sculpture associated with posterior meso-
pleural furrow. — Fischer (1972) segregated
species in Opius into different subgenera
based on the presence or absence of cren-
ulations within the posterior mesopleural
furrow. This interpretation is inadequate
because within a species the posterior me-
sopleural furrow can be either entirely
smooth or crenulate below but smooth
above the episternal scrobe. Nearctic spe-
cies in Gastrosema that exhibit this condi-
tion include O. abbyae Kula, new species
(described below) (Figs. 15-16).

An entirely smooth posterior meso-
pleural furrow is found in the majority of
Nearctic species in Gastrosema. Yet several
species in Gastrosema consistently possess

Volume 12, Number 2, 2003

283

Figs. 15-16. Posterior mesopleural furrow. 15, Entirely smooth. 16, Crenulate below (arrow a) but smooth
above (arrow b) episternal scrobe. Figs, modified from Fischer (1972).

an entirely crenulate posterior mesopleur-
al furrow (Fig. 1). Opius flaviceps, O. rela-
tivus, O. salmonensis, and O. tangens are
Nearctic species in Gastrosema that possess
an entirely crenulate posterior mesopleur-
al furrow. Additionally, several Nearctic
species in Pleurosema consistently possess
an entirely crenulate posterior mesopleur-
al furrow. When interpreting this charac-
ter in phylogenetic analyses, the location
of crenulations in the posterior mesopleur-
al furrow is more informative than the
mere presence or absence of crenulations
and may be important for establishing ho-
mology.

Presence or absence of mesoscntal midpit. —
Fischer (1972) used the presence or ab-
sence of the mesoscutal midpit (Fig. 17) to
divide the subgenera of Opius into two

roughly equal sized groups. The assess-
ment of variation conducted in this study
confirms the potential phylogenetic utility
of this character. Midpit shape and depth
vary intraspecifically in Opiinae, but the
midpit is consistently present or absent for
a particular species. All Nearctic species in
Gastrosema consistently lack a midpit, but
there are several Nearctic species in other
subgenera that consistently possess a mid-
pit. Because the midpit can be quite re-
duced within a species, specimens should
be examined at the appropriate angle and
with light dispersing plastic (mylar).

Presence, absence, and sculpture associated
with notauli. — Species in Opiinae exhibit
several character states associated with the
notauli. Historically, the length of the no-
tauli has been used to group putatively

Figs. 17-18. 17, Mesoscutal midpit. 18, Notauli represented by shallow anterior depressions.

284

Journal of Hymenoptera Research

Figs. 19-20. Sternaulus. 19, Short, narrow, and crenulate. 20, Represented by a smooth depression.

closely related species (Fischer 1972).
However, the presence or absence of cren-
ulations within the notauli, as well as the
termination point of the notauli, may pro-
vide additional useful information for
phylogenetic inference. Among Nearctic
species in Gastrosema, the notauli are shal-
low or deep anterior depressions that can
be sculptured or unsculptured (Fig. 18).

Presence, absence, and sculpture associated
with sternaulus. — Fischer (1972) used the
following three attributes to partially de-
fine certain subgenera: sternaulus absent,
sternaulus present and smooth, and ster-
naulus present and sculptured (e.g. cren-
ulate). Gastroscma is partially defined by
the presence of sculpture within the ster-
naulus. The most commonly observed
condition for Nearctic species in Gastrose-
ma is a relatively short and narrow ster-
naulus with crenulate sculpture (Fig. 19).
However, examination of multiple indi-
viduals for several Nearctic species in Gas-
troscma revealed that the sternaulus can
vary from a smooth depression (Fig. 20) to
distinctly crenulate within a species. Spe-
cies that exhibit this condition include O.
lacopitaensis and O. striativentris. Further,
species such as O. castaneigaster Fischer of-
ten possess a very weakly sculptured ster-
naulus that can appear unsculptured us-
ing a stereomicroscope with a fiber optic
illuminator. Despite this large breadth of
variation, Fischer (1972) used the presence

and absence of sculpture within the ster-
naulus to place species in Gastrosema and
Phaedrotoma, respectively. Because the
presence and absence of sculpture within
the sternaulus can vary intraspecifically,
certain Nearctic species in Phaedrotoma
sensu Fischer (1972) potentially share a
most recent common ancestor with certain
Nearctic species in Gastrosema. The high
level of intraspecific variation in sternau-
lus sculpture limits the use of this char-
acter in phylogenetic analyses. If sculpture
within the sternaulus is utilized in future
phylogenetic analyses, the location of the
sternaulus and the type of sculpture with-
in the sternaulus are important for estab-
lishing homology.

Sculpture associated with propodeum. —
The presence or absence of propodeal
sculpture is a major defining feature for
several subgenera in Opius. In Fischer
(1972) the presence or absence of propo-
deal sculpture is the only attribute differ-
entiating species in Merotrachys and Phae-
drotoma, respectively. The presence, ab-
sence, and type of propodeal sculpture are
quite variable among Nearctic species in
Gastrosema, but these attributes are only
slightly variable within a species. For ex-
ample, in O. striativentris the propodeum
is rugulose to rugose (Fig. 21), in O. inter-
missus Fischer the propodeum is always
rugose, and in O. salmonensis the propo-
deum is always areolate-rugose (Fig. 22).

Volume 12, Number 2, 2003

285

Figs. 21-23. Propodeal sculpture. 21, Rugulose. 22, Areolate-rugose. 23, Smooth with rugulose sculpture
surrounding spiracles.

The majority of Nearctic species in Gastro-
sema possess a rugulose to rugose propo-
deum. However, species that possess an
areolate-rugose propodeum are also com-
monly encountered. Further, species with
an areolate-rugose propodeum usually
have an entirely crenulate posterior me-
sopleural furrow. Nearctic species that ex-
hibit this condition include O. flaviceps, O.
relativus, O. salmonensis, and O. tangens.
The majority of Nearctic species in Pleu-
rosema also possess an areolate-rugose
propodeum and an entirely crenulate pos-
terior mesopleural furrow. Thus, certain
Nearctic species in Pleurosema potentially
share a most recent common ancestor with
certain Nearctic species in Gastrosema. Of
the described Nearctic species in Gastro-
sema, only O. haucockamis Fischer has a
nearly entirely smooth propodeum (Fig.

23). A trace of rugulose sculpturing sur-
rounds the propodeal spiracles, but the
rest of the propodeum is completely
smooth. This condition is also commonly
observed for Nearctic species in Phaedro-
toma sensu Fischer (1972).

Propodeal sculpture can be assigned
distinct character states, and different
types of propodeal sculpture often corre-
late with sculpture in the posterior meso-
pleural furrow. Species with an areolate-
rugose propodeum frequently have an en-
tirely crenulate posterior mesopleural fur-
row, and species with a smooth or
rugulose to rugose propodeum typically
do not have an entirely crenulate posterior
mesopleural furrow. Consideration of pro-
podeal sculpture in phylogenetic analyses
must go beyond the presence or absence
of sculpture. Different types of propodeal

286

Journal of Hymenoptera Research

sculpture, and the correlation between
propodeal sculpture and sculpture asso-
ciated with other anatomical features,
should be considered.

Sculpture associated with median abdomi-
nal tergites. — In Fischer's original diagno-
sis for Gastrosema, a major defining feature
was the presence of "shagreened" or co-
riaceous sculpture on abdominal tergite
two (t2) (Fig. 24) (Fischer 1972). Exami-
nation of the Nearctic species in Gastrose-
ma treated by Fischer (1977) revealed that
Gastrosema was not limited to species with
a coriaceous t2. Rather, species with a car-
inate or costate t2 were included as well
(Figs. 25-26). The vast majority of Nearctic
species included in Gastrosema have a co-
riaceous t2. Examples include O. abbyae, O.
castaneigaster, O. intermissus, O. lacopitaen-
sis, and O. striativentris. Opius flaviceps and
O. salmonensis are two species that possess
a costate t2, and O. oscinidis possesses a
carinate t2. Neither costate nor carinate
sculpture on t2 is common among Nearc-
tic species in Gastrosema, but a carinate t2
is commonly observed for several Nearctic
species in Rhogadopsis sensu Wharton
(1987) and Thoracosema. In terms of intra-
specific variation, species with a coria-
ceous t2 show little variation in sculpture
density. Conversely, species with a cari-
nate or costate t2 exhibit substantial intra-
specific variation in sculpture density. In
O. oscinidis the carinations are extremely
reduced in some individuals, and t2 ap-
pears smooth unless the specimens are ex-
amined at several angles using mylar and
a variety of light intensities. Yet the cari-
nations on t2 are unmistakably present in
other individuals. Thus, sculpture on t2
has potential phylogenetic importance but
may be difficult to code unambiguously
for certain species.

Configuration of forewing venation. — Fi-
scher (1972) used the position of vein lm-
cu relative to vein 2RS to segregate species
into certain subgenera. For example, spe-
cies that Fischer placed in Phlebosema Fi-
scher possess vein lm-cu basad or directly

in line with vein 2RS (Figs. 27-28). Phle-
bosema is currently a junior synonym of
Baeocentrum Schulz, but this nomenclatur-
al change does not alter Fischer's concept
of Phlebosema (Wharton 1987). Alternative-
ly, species that Fischer placed in Gastrose-
ma possess vein lm-cu distad vein 2RS
(Fig. 29). The results of this study confirm
that species with vein lm-cu basad vein
2RS occasionally display vein lm-cu di-
rectly in line with vein 2RS. Further, spe-
cies with vein lm-cu distad vein 2RS nev-
er exhibit vein lm-cu directly in line with
or basad vein 2RS. All described Nearctic
species in Gastrosema have vein lm-cu dis-
tad vein 2RS. However, the Nearctic spe-
cies in Baeocentrum sensu Wharton (1987)
and certain Nearctic species in Merotrachys
and Tolbia sensu Fischer (1972) have vein
lm-cu basad or directly in line with vein
2RS. Thus, the position of vein lm-cu rel-
ative to vein 2RS should be evaluated in
future phylogenetic analyses.

The size and shape of the forewing stig-
ma is another character of potential phy-
logenetic importance. The Nearctic species
in Gastrosema exhibit minimal intraspecific
variation in stigma size and shape, but
stigma size and shape can vary substan-
tially from species to species. Several Ne-
arctic species in Gastrosema exhibit con-
spicuous similarities in stigma size and
shape. Opius abbyae, O. castaneigaster, O.
intermissus, O. lacopitaensis, and O. striativ-
entris possess a narrow and elongate stig-
ma (Fig. 28), while O. flaviceps, O. relativus,
O. salmonensis, and O. tangens possess a
wedge shaped stigma (Fig. 29). In future
studies stigma size and shape should be
thoroughly examined for species in other
opiine subgenera.

Color. — In Opiinae color is not of any
apparent phylogenetic importance but is
useful for distinguishing between mor-
phologically similar species. For example,
O. intermissus and O. niobe Fischer are sim-
ilar in size and shape but have conspicu-
ously different coloration on the head and
mesosoma. Species such as O. striativentris

Volume 12, Number 2, 2003

287

Figs. 24-26. Sculpture on t2. 24, Coriaceous. 25, Carinate. 26, Costate.

are variable in color, while color is more
conserved in species such as O. abbyae.
Thus, it is important to examine several
individuals to determine the entire spec-
trum of color within a species. Proper
specimen preparation is essential for pre-
serving natural coloration. Specimens ex-
posed to light for prolonged periods of
time become lighter, and air-dried speci-
mens tend to darken. Material preserved
in alcohol for extended periods of time
should be stored at low temperatures and
protected from direct exposure to light.
Specimens should be dehydrated using a
critical-point-dryer or chemicals such as
amyl acetate and Hexamethyl-Disilazane.
Setation. — Setation is another useful
character for distinguishing between mor-
phologically similar species. For example,
a species of Opius near extiratus Fischer
possesses a consistently setose propo-

deum, while setation on the propodeum
of O. striativentris is usually setiferous and
rarely glabrous. Opius near extiratus and
O. striativentris are sympatric and have
been reared from the same host species on
the same host plant. The density of setae
on the propodeum is one character that
can be used to distinguish these species.

Flagellomere number. — Godfray (1984)
demonstrated that in the alysiine Exotela
cyclogaster Forster flagellomere number in-
creases with body size. Alysiinae and Opi-
inae form a monophyletic group (Wharton
1988), and it is possible that species in
Opiinae also exhibit this trend. Flagello-
mere number is useful when morpholog-
ically similar species have non-overlap-
ping ranges of flagellomere number. For
example, O. abbyae and O. intennissus are
morphologically similar, but O. abbyae has
19-22 flagellomeres compared to 26-29 in

288

Journal of Hymenoptera Research

27

28

29

Figs. 27-29. Position of forewing vein lm-cu (arrow
a) relative to vein 2RS (arrow b). 27, lm-cu basad 2RS.
28, lm-cu directly in line with 2RS. 29, lm-cu distad
2RS.

O. intermissus. When using flagellomere
number to distinguish between species, it
is important to examine several individu-
als to establish minimum and maximum
flagellomere numbers for each species.

Quantitative characters. — Fischer (1972,
1977, 1987) extensively used quantitative
ratios in keys to subgenera, species
groups, and species in Opius, but the ra-
tios were based on measurements of one
or a few individuals. As a result most taxa
treated in the keys cannot be reliably iden-
tified. A prime example is the treatment of
species groups in Fischer's (1977) key to
New World Gastrosema. The defining char-
acter of the lissopleurum-group of species
is "thorax 1.25-1. 40 X as long as high."
Opius intermissus is included in the lisso-

pleurum-group, but the results of this
study indicate that the thorax is 1.12-
1.30X as long as high in O. intermissus.
Only 33.0% of the specimens examined for
O. intermissus fit within the morphological
limits of the lissopleurum-group as defined
by Fischer (1977).

Another problem with the use of quan-
titative ratios is that measurement error as
high as 15.7% can occur when the same
specimen is measured on different dates.
Factors that contribute to measurement er-
ror may be found in Wharton (1980). Giv-
en the limitations of quantitative char-
acters, it is preferable to use qualitative
characters in phylogenetic analyses and
identification systems. However, measure-
ments can be useful for distinguishing cer-
tain species. Opius flaviceps and O. salmo-
nensis are similar in terms of coloration
and sculpture, but the two species can be
differentiated using ovipositor length, me-
sosoma length, and mesosoma width.
When using measurements to characterize
a species, it is optimal to measure several
individuals to establish the entire range of
variation for each measurement. In cases
where only one or a few individuals are
available for examination, the taxonomist
should consider the usefulness of quanti-
tatively describing the species.

NEARCTIC SPECIES IN THE
SUBGENUS GASTROSEMA

Species group diagnoses. — With the excep-
tion of O. walleyi, all described Nearctic
species in Gastrosema fall into three easily
characterized groups that have not been
recognized in previous studies (Table 1).
A diagnosis of each group is provided to
facilitate future studies examining the
monophyly and relationships of species in
Opius. Each group should be viewed as a
hypothesis supported by potentially phy-
logenetically informative characters, and
the monophyly of each group will be test-
ed in future phylogenetic analyses. The
first three characters presented in each di-

Volume 12, Number 2, 2003

289

Table 1. Placement of described Nearctic species
in Gastrosema into three previously unrecognized spe-
cies groups.

flatriceps-

group

oscinidis-
group

striativentris-

group

flaviceps
relativus
salmonensis

oscinidis

abbyae
adductus
alconanus Fischer

tan gens

castaneigaster

clevelandeiisis

cordobensis

deiphobe Fischer

liaiicockaiius

intermissus

lacopitaensis

niobe

pallas

porteri Fischer

striativciitris

agnosis differentiate each group from the
remaining Opius sensu lato.

flaviceps-group: midpit absent; clypeus
reduced, labrum exposed when mandibles
closed; lm-cu distad 2RS; posterior me-
sopleural furrow crenulate above epister-
nal scrobe; propodeum rugose to areolate-
rugose; abdominal tergites posterior to tl
smooth or variously sculptured; forewing
stigma usually broad and wedge shaped,
second submarginal cell size and shape
variable but often shaped as in Fig. 29.

oscinidis-group: midpit absent; clypeus
reduced, labrum exposed when mandibles
closed; lm-cu distad 2RS; posterior me-
sopleural furrow smooth above episternal
scrobe; propodeum smooth to rugose; ab-
dominal tergites posterior to tl smooth or
longitudinally carinate; forewing stigma
usually narrow and elongate, second sub-
marginal cell long and usually shaped as
in Fig. 30.

striativentris-group: midpit absent; clyp-
eus reduced, labrum exposed when man-
dibles closed; lm-cu distad 2RS; posterior
mesopleural furrow smooth above epister-
nal scrobe; propodeum smooth to rugose;
at least one abdominal tergite posterior to
tl coriaceous; forewing stigma usually

Figs. 30-31. Shape of forewing stigma and second
submarginal cell. 30, oscinidis-group. 31, striativentris-
group.

narrow and elongate, second submarginal
cell short and usually shaped as in Fig. 31.

Discussion. — The posterior mesopleural
furrow in O. walleyi is smooth above the
episternal scrobe, and thus, O. walleyi does
not fall within the flaviceps-group. The
shape of the second submarginal cell in O.
walleyi fits the diagnosis for the oscinidis-
group. However, t2 and t3 are coriaceous
which fits the diagnosis for the striativen-
tris-group. Thus, Opius walleyi possesses
characteristics of both the oscinid /s-group
and the striativentris-group and is left un-
placed at this time.

Opius pumilio (the type species for Gas-
trosema) belongs in the striativentris-group.
Additionally, several Nearctic species
treated in Fischer (1977) were not included
in Gastrosema but belong in one of the
three species groups (Table 2). Thus, if the
species groups proposed in this study are
monophyletic, subgenera with species that
fall within the species groups are obvious-
ly not monophyletic.

In future studies of Opius, species in Al-
lophlebus Fischer, Gastrosema, Hypocynodus
Forster sensu Fischer (1972), Merotrachys,
Opiothorax Fischer, Peudopius Fischer,
Phaedrotoma sensu Fischer (1972), Pleurose-

290

Journal of Hymenoptera Research

Table 2. Placement of described Nearctic species not included in Gastrosema into three previously unrec-
ognized species groups.

flaviceps-group

O. (Pleurosema)
euwattacooanus Fischer
hermosanus Fischer
metatensis Fischer
orizabensis Fischer
paratakomanus Fischer
pilosinotum Fischer
sybile Fischer
thalis Fischer

O. (Rhogadopis)
northcarolinensis Fischer

oscinidis-gioxip

O. (Rhogadopis)

stenopectus Fischer
O. (Thoracosema )

crabtreeanus Fischer

extiratus

parkercrcekensis Fischer

prolongatus Fischer

pseudocolumbiacus Fischer

schuleri Fischer

southcarolinensis Fischer

striativentris-group

O. (Merotrachys)

brownsvillensis Fischer

lansingensis Fischer

paulior Fischer

paulus Fischer

virentis Fischer
O. (Phaedrotoma)

complicans Fischer
O. (Tolbia)

heroicus Fischer

ma, Rhogadopisis sensu Wharton (1987),
Thoracosema, and Tolbia sensu Fischer
(1972) should be examined to determine
which species group, if any, they fit into.
Examination of species in the aforemen-
tioned subgenera will almost certainly re-
sult in the discovery of species groups in
addition to the three discovered during
this research. Placement of all described
species in these subgenera will be a mon-
umental task, as several hundred holo-
types must be examined. The holotypes
are housed in repositories found through-
out the world, although most are in North
American and European museums and
collections. Species currently included in
subgenera not listed above lack at least
one of the three characters used to differ-

entiate the three species groups from the
remaining Opius sensu lato (see species
group diagnoses).

Opius (Gastrosema) abbyae Kula,
new species

Fig. 32

Female. — Head: 1. 23-1. 36 X as wide as
long; eyes 0.67-0.77X as wide as long, gla-
brous; face smooth with rugulose sculp-
ture between antennal sockets and com-
pound eyes, slightly raised mesally, seti-
ferous; clypeus hemielliptical, convex,
ventral margin straight, setiferous; man-
dibles simple, with two apical teeth per
mandible, setiferous; antennae with 19-21
flagellomeres. Mesosoma: 0.30-0.40 mm
wide; 0.51-0.64 mm long; 0.58-0.66 X as

Figs. 32-33. Opius spp. 32, O. abbyae. 33, O. lacopitaensis.

Volume 12, Number 2, 2003

291

wide as long; 0.71-0.81 X as wide as deep;
1.10-1.31 X as long as deep; pronotum co-
riaceous laterally, crenulate in anterior
furrow; notauli represented by anterior
depressions; mesoscutum smooth, setifer-
ous (78.6%) or sparsely setiferous anteri-
orly and along lines where notauli would
run if present (21.4%); mesopleuron nearly
entirely smooth, usually rugulose or cren-
ulate near tegulae (85.8%), rarely smooth
near tegulae (14.2%), sternaulus distinctly
crenulate; scutellar disc smooth; propo-
deum rugose, setiferous; metapleuron se-
tiferous. Metasoma: tl 0.93-1.25 X as wide
as long, rugulose to rugose; t2 coriaceous;
t3 coriaceous; t4 always at least partially
coriaceous; remaining tergites smooth; ex-
posed ovipositor 0.11-0.26 mm long. Legs:
smooth, setiferous, hind tibia 0.95-1. 09 X
as long as mesosoma length. Wings: hya-
line; forewing stigma narrow and elon-
gate; 2RS 0.53-0.71 X as long as 3RSa; 2RS
0.41-0.50X as long as 2M; 3RSa 0.70-0.77X
as long as 2M. Color: head entirely very
dark brown to black; mesosoma usually
black (85.7%), rarely very dark brown
(14.3%); tl entirely dark brown to nearly
entirely dark brown with slight brownish
orange coloration laterally; t2 light brown
to dark brown medially, orange to yellow
laterally; t3 usually brown to dark brown
with orange to yellow coloration at an-
terolateral edges (85.7%), rarely entirely
dark brown (14.3%); t4 usually entirely
dark brown (92.9%), rarely brownish yel-
low (7.1%); remaining tergites dark
brown; legs yellow to yellowish brown,
prothoracic legs lighter than mesothoracic
legs, mesothoracic legs lighter than meta-
thoracic legs, metatibia and metatarsus al-
ways yellowish brown.

Male.— As in 9 except: Head: 1.21-1. 30 X
as wide as long; eyes 0.64-0.71 X as wide
as long; antennae with 20-22 flagello-
meres. Mesosoma: 0.57-0.61 X as wide as
long; 1.24-1. 32 X as long as deep. Wings:
2RS 0.50-0.53 X as long as 2M. Color: me-
sosoma always black.

Host. — Calycomyza sp. mining the leaves
of Helianthus annuus (TAMU).

Material examined. — Holotype 9: USA,
Nebraska, Arthur County, Highway 61
south of Arthur near Arapaho Prairie,
8.VU.1999, R. R. Kula and A. A. Rogers
(TAMU). Paratypes: 69, same data as ho-
lotype; 29, Id, same data as holotype ex-
cept ll.vii.1998; 59, 3d, same data as ho-
lotype except 10.vii.1999 (TAMU).

Diagnosis. — Within the striativentris-
group O. intermissus, O. lacopitaensis, and
O. weemsi Fischer are morphologically
similar to O. abbyae. Head coloration can
be used to distinguish abbyae from both in-
termissus and weemsi. In abbyae the head is
entirely very dark brown to black. Alter-
natively, in intermissus the head is yellow
with the vertex very dark brown to black,
and in weemsi the head is mostly orange
with the vertex very dark brown and the
ventral half of the face brown. The num-
ber of antennal flagellomeres can also be
used to distinguish abbyae from intermissus
and weemsi. Opius abbyae possesses 19-22
flagellomeres, while intermissus possesses
26-29 flagellomeres and weemsi possesses
32 flagellomeres. Another character that
can be used to differentiate abbyae from in-
termissus and weemsi is tergite coloration.
Tergite 3 in abbyae is usually brown to
dark brown with orange to yellow color-
ation at the antero-lateral edges. In inter-
missus t3 is yellow with the posterior edge
dark brown, and in weemsi t3 is orangish
yellow with the posterior edge dark
brown. Sculpture on the abdominal ter-
gites can be used to distinguish abbyae
from lacopitaensis. In abbyae t4 is always co-
riaceous, while in lacopitaensis t4 is never
coriaceous.

Discussion. — Opius abbyae falls within
the striativentris-group. This species exhib-
its less intraspecific variation in qualitative
characters than observed for intermissus,
lacopitaensis, oscinidis, and striativentris. For
example, the sternaulus is consistently
crenulate in abbyae, while the sternaulus
varies from a smooth depression to cren-

292 Journal of Hymenoptera Research

ulate in lacopitaensis and striativentris. Col- Wings: hyaline; forewing stigma narrow

oration of the head and mesosoma is and elongate; 2RS 0.56-0.64X as long as

much less variable in abbyae than in oscin- 3RSa; 2RS 0.41-0.48X as long as 2M; 3RSa

idis or striativentris. No substantial differ- 0.67-0.77X as long as 2M. Color: head en-

ences were observed between female and tirely dark brown to very dark brown; me-

male specimens in abbyae. sosoma entirely dark brown to very dark

Etymology. — This species is named in brown; tl dark brown; t2 brownish orange

honor of Abigail Rogers Kula who assist- to yellow; t3 anterior half to two thirds

ed in collection of the host. brownish orange to yellow, posterior half

to one third brown; remaining tergites

Opins (Gastrosema) lacopitaensis Kula, brown to dark brown; legs yellow, poste-

new species rjor p0rnon of metatibia and entire meta-

Fig- 33 tarsus yellowish brown.

Female.— Head: 1.25-1. 36 X as wide as Male.— As in 9 except: Head: eyes 0.67-

long; eyes 0.67-0.73 X as wide as long, gla- 0.79 X as wide as long; face usually

brous; face usually smooth with rugulose smooth with rugulose sculpture between

sculpture between antennal sockets and antennal sockets and compound eyes

compound eyes (92.9%), rarely minutely (90.9%), rarely entirely smooth (9.1%); an-

coriaceous with rugulose sculpture be- tennae with 19-22 flagellomeres. Mesoso-

tween antennal sockets and compound ma: 0.54-0.67 mm long; 0.68-0.73 X as

eyes (7.1%), slightly raised mesally, setifer- wide as deep; 1.17-1.38X as long as deep;

ous; clypeus narrowly hemielliptical to pronotum often coriaceous laterally

crescent shaped, convex, ventral margin (63.6%), occasionally smooth (36.4%); me-

straight to slightly arched, setiferous; man- sopleuron nearly entirely smooth, usually

dibles simple to gradually widened distally smooth near tegulae (90.9%), rarely cren-

to proximally, with two apical teeth per ulate near tegulae (9.1%), sternaulus often

mandible, setiferous; antennae with 19-20 a smooth depression (54.6%), occasionally

flagellomeres. Mesosoma: 0.29-0.35 mm crenulate (45.5%). Wings: 2RS 0.56-0.67X

wide; 0.51-0.61 mm long; 0.55-0.64X as as long as 3RSa.

wide as long; 0.69-0.75 X as wide as deep; Host. — Unknown.

1.14-1.27X as long as deep; pronotum of- Material examined. — Holotype 9: USA,
ten entirely coriaceous laterally (78.6%), oc- Texas, Jim Wells County, 8 miles west of
casionally smooth dorsally and coriaceous Ben Bolt, La Copita Research Station,
ventrally (21.4%), crenulate in anterior fur- 20.V.1987, J. B. Woolley (TAMU). Para-
row; notauli represented by anterior de- types: 11$, 11<3, same data as holotype;
pressions; mesoscutum smooth, setiferous 2$, same data as holotype except north-
anteriorly and along lines where notauli west of Ben Bolt, 21. v. 1987, G. Zolnerow-
would run if present; mesopleuron nearly ich (TAMU).

entirely smooth, often smooth near tegulae Diagnosis. — Within the striativentris-

(78.6%), occasionally crenulate near tegulae group O. abbyae, O. castaneigaster, O. cor-

(21.4%), sternaulus often crenulate (78.6%), dobensis Fischer, and O. pallas Fischer are

occasionally a smooth depression (21.4%); morphologically similar to O. lacopitaensis.

scutellar disc smooth; propodeum rugose, Characters used to differentiate lacopitaen-

setiferous; metapleuron setiferous. Metaso- sis from abbyae and castaneigaster can be

ma: tl 0.71-1.00X as wide as long, rugose; found in the diagnoses for abbyae and cas-

t2 coriaceous; t3 coriaceous; remaining ter- taneigaster. Optus lacopitaensis can be dif-

gites smooth; exposed ovipositor 0.16-0.32 ferentiated from cordobensis using propo-

mm long. Legs: smooth, setiferous, hind tib- deal sculpture. In lacopitaensis the propo-

ia 0.88-0.97X as long as mesosoma length, deum is rugulose to rugose, while in cor-

Volume 12, Number 2, 2003

293

dobensis the propodeum is very heavily
rugose with conspicuous carinae through-
out. The number of antennal flagellomeres
can be used to distinguish lacopitaensis
from pallas. Optus lacopitaensis possesses
19-22 flagellomeres, while pallas possesses
28 flagellomeres.

Discussion. — O. lacopitaensis falls within
the striativentris-group. In Fischer's sub-
generic classification individuals with a
crenulate sternaulus fall within Gastrose-
ma, but individuals with a smooth ster-
naulus fall within Merotrachys. Opius lacop-
itaensis is provisionally placed in Gastro-
sema because the majority of individuals
(i.e. 16 of 25) fall within Gastrosema.

Opius lacopitaensis exhibits variation in
color and sculpture similar to that ob-
served in castaneigaster. No substantial dif-
ferences were observed between female
and male specimens in lacopitaensis. No bi-
ological data are associated with lacopi-
taensis, but other species in the striativen-
t r/s-group have been reared from leaf min-
ing Agromyzidae (e.g. O. abbyae, O. adduc-
tus Fischer, O. striativentris).

Etymology. — This species is named after
the collection site.

Opius (Gastrosema) castaneigaster
Fischer

Opius castaneigaster Fischer 1964: 413, 419.
Opius (Gastrosema) castaneigaster: Fischer 1977:
311, 324.

Female. — Head: 1.19-1. 33 X as wide as
long; eyes 0.62-0.75 X as wide as long, gla-
brous; face usually smooth with rugulose
sculpture between antennal sockets and
compound eyes (91.7%), rarely entirely
smooth (8.3%), slightly raised mesally, se-
tiferous; clypeus hemispherical, convex,
ventral margin straight, setiferous; man-
dibles simple, with two apical teeth per
mandible, setiferous; antennae with 21-24
flagellomeres. Mesosoma: 0.30-0.45 mm
wide; 0.54-0.82 mm long; 0.53-0.64 X as
wide as long; 0.79-0.93 X as wide as deep;
1.29-1. 70 X as long as deep; pronotum co-

riaceous laterally, crenulate in anterior
furrow; notauli represented by anterior
depressions; mesoscutum smooth, setifer-
ous anteriorly and along lines where no-
tauli would run if present; mesopleuron
nearly entirely smooth, usually rugulose
or crenulate near tegulae (91.7%), rarely
smooth near tegulae (8.3%), sternaulus
usually crenulate (91.7%), rarely reduced
to a sculptured depression (8.3%); scutel-
lar disc smooth; propodeum rugulose to
rugose, setiferous; metapleuron sparsely
setiferous to setiferous. Metasoma: tl 1.00-
1.33 X as wide as long, rugulose to rugose;
t2 usually coriaceous (91.7%), rarely
smooth (8.3%); t3 usually at least partially
coriaceous (91.7%), rarely entirely smooth
(8.3%); remaining tergites smooth; ex-
posed ovipositor 0.24-0.49 mm long. Legs:
smooth, setiferous, hind tibia 0.83-0.95 X
as long as mesosoma length. Wings: hya-
line; forewing stigma narrow and elon-
gate; 2RS 0.48-0.67X as long as 3RSa; 2RS
0.39-0.50X as long as 2M; 3RSa 0.73-0.84X
as long as 2M. Color: head entirely brown
to dark brown; mesosoma entirely brown
to dark brown; tl brown to dark brown;
t2 orange to yellow; t3 anterior half orange
to yellow, posterior half brown to dark
brown or t3 anterior two thirds orange to
yellow, posterior one third brown to dark
brown; remaining tergites dark brown;
legs orangish yellow to yellow, metatibia
and metatarsus usually yellowish brown
(83.3%).

Male.— As in 9 except: Head: 1.1 1-1. 26 X
as wide as long. Mesosoma: 0.51-0.60X as
wide as long. Metasoma: tl 0.86-1. 17X as
wide as long.

Host. — Unknown.

Material examined. — Holotype 9: USA,
New York, Tompkins County, Ithaca,
7.ix.l935, H. K. Townes (AEI). Allotype 6:
same data as holotype except l.vi.1935
(AEI). Paratypes: all USA; 19, same data
as holotype except 30. v. 1934; 16, New
York, Oneida County, Rome, 24.vi.1934,
H. K. Townes; 19, New York, Delaware
County, Hancock, 31.vii.1935, H. K.

294

Journal of Hymenoptera Research

Townes; 1 9 , 1 6, New York, Otsego Coun-
ty, Oneonta, 17.viii.1935, H. K. Townes;
19, New York, Otsego County, Oneonta
swamp, 1,900 feet elevation, 18.viii.1935,
H. K. Townes; 1 9 , Ohio, Summit County,
Akron, 18.V.1941, H. K. Townes; 1 9, South
Carolina, Greenville County, Greenville,
22.iv.1952, G. and L. Townes; 19, South
Carolina, Greenville County, Greenville,
31.V.1952, G. and L. Townes; 19, South
Carolina, Pickens County, Wattacoo,
lO.v.1959, G. F. Townes; 16, South Caro-
lina, Pickens County, Wattacoo, 6. v. 1961,
G. F. Townes; 1 9 , South Carolina, Pickens
County, Wattacoo, 19. v. 1961, G. F.
Townes (AEI); 1 9 , Wisconsin, Polk Coun-
ty, July, collection C. F. Baker (USNM).
Other determined material: 1 9 , same data
as holotype (det. Fischer) (AEI).

Diagnosis. — Within the striativentris-
group, O. cordobensis, O. lacopitaensis, O.
niobe, O. pallas, and O. striativentris are
morphologically similar to O. castaneigas-
ter. Propodeal sculpture can be used to
distinguish castaneigaster from cordobensis.
In castaneigaster the propodeum is rugu-
lose to rugose, while in cordobensis the pro-
podeum is very heavily rugose with con-
spicuous carinae throughout. Opius casta-
neigaster can be differentiated from niobe
on the basis of mesosoma shape and the
number of antennal flagellomeres. In cas-
taneigaster the mesosoma is subelliptical
(1.29-1. 70 X as long as deep), and the
number of flagellomeres ranges from 21-
24. In niobe the mesosoma is subspherical
(1.21-1.23 X as long as deep), and the
number of flagellomeres ranges from 25-
31. Opius pallets is very morphologically
similar to castaneigaster, but the two spe-
cies can be differentiated using flagello-
mere number. Opius pallas possesses 28
flagellomeres, while the maximum num-
ber of flagellomeres for castaneigaster is 24.
Several characters can be used to distin-
guish castaneigaster from striativentris. Ter-
gite 1 is brown to dark brown in castanei-
gaster, while tl is usually yellow to brown-
ish orange in striativentris. Tergite 3 is only

partially coriaceous in castaneigaster, as op-
posed to entirely coriaceous in striativen-
tris. Tergite 4 is always smooth in casta-
neigaster, while t4 is smooth or coriaceous
in striativentris. The mesosoma is uniform-
ly brown or dark brown in castaneigaster,
while the mesonotum is often conspicu-
ously darker than the rest of the mesoso-
ma in striativentris. Opiits castaneigaster can
be differentiated from lacopitaensis using tl
width to length ratios, clypeus shape, and
the number of flagellomeres. Tergite 1
width in castaneigaster is usually greater
than or equal to tl length (91.7%) and is
rarely less than tl length (8.3%). Tergite 1
width in lacopitaensis is always less than or
equal to tl length. The shape of the clyp-
eus is hemispherical in castaneigaster,
while the shape of the clypeus is narrowly
hemielliptical to crescent shaped in lacop-
itaensis. Opiits castaneigaster possesses 21-
24 flagellomeres, as opposed 19-22 in la-
copitaensis.

Discussion. — Opius castaneigaster falls
within the striativentris-group. Opius pmllas
may eventually be determined as a syno-
nym of castaneigaster. The two species are
very morphologically similar and exhibit
overlapping geographic distributions. Op-
ius pallas is only known from the holotype,
and acquisition of additional specimens
may reveal a broader range of flagello-
mere number.

One specimen of castaneigaster exam-
ined in this study lacks sculpture on t2
and t3. However, no additional attributes
were observed that indicated the speci-
men was not castaneigaster. Single speci-
mens with reduced sculpture are very dif-
ficult to identify, and comparison of these
individuals with a long series of deter-
mined specimens will facilitate their iden-
tification. No biological data are associat-
ed with castaneigaster, but other species in
the striativentris-group have been reared
from leaf mining Agromyzidae (e.g. O. ab-
byae, O. adductus, O. striativentris).

Volume 12, Number 2, 2003

295

Opius (Gastrosema) intermissus Fischer

Opius intermissus Fischer 1964: 413, 428.
Opius {Gastrosema) intermissus: Fischer 1977:
314, 357.

Female. — Head: 1.23-1.29 X as wide as
long; eyes 0.62-0.67X as wide as long, gla-
brous; face minutely coriaceous with ru-
gulose sculpture between antennal sockets
and compound eyes, slightly raised mes-
ally, usually setose (80.0%), rarely setifer-
ous (20.0%); clypeus hemispherical, con-
vex, ventral margin straight, usually se-
tose (80.0%), rarely setiferous (20.0%);
mandibles simple, with two apical teeth
per mandible, usually setose (80.0%), rare-
ly setiferous (20.0%); antennae with 27-29
flagellomeres. Mesosoma: 0.38-0.46 mm
wide; 0.64-0.75 mm long; 0.59-0.63 X as
wide as long; 0.71-0.74 X as wide as deep;
1.14-1.21 X as long as deep; pronotum co-
riaceous laterally, crenulate in anterior
furrow; notauli represented by anterior
depressions; mesoscutum smooth, setifer-
ous anteriorly and along lines where no-
tauli would run if present; mesopleuron
nearly entirely smooth, rugulose to cren-
ulate near tegulae, sternaulus distinctly
crenulate; scutellar disc smooth; propo-
deum rugose, setiferous; metapleuron
usually setiferous (80.0%), rarely setose
(20.0%). Metasoma: tl 0.70-1. 20 X as wide
as long, longitudinally rugulose with two
prominent dorsal carinae converging pos-
teriorly; t2 coriaceous; t3 coriaceous; re-
maining tergites smooth; exposed ovipos-
itor 0.18-0.32 mm long. Legs: smooth, se-
tiferous, hind tibia 0.96-1. 02 X as long as
mesosoma length. Wings: hyaline; fore-
wing stigma narrow and elongate; 2RS
0.52-0.62 X as long as 3RSa; 2RS 0.41-
0.47X as long as 2M; 3RSa 0.75-0.82X as
long as 2M. Color: head mostly yellow,
vertex very dark brown to black; meso-
soma very dark brown to black, meso-
pleuron occasionally yellowish orange
ventrally and near tegulae; tl brown to
dark brown medially, yellow laterally; t2
yellow; t3 yellow with posterior edge dark

brown; t4 dark brown medially, yellow
laterally, and posterior edge dark brown
(60.0%) or t4 mostly yellow with posterior
edge dark brown (40.0%); remaining ter-
gites dark brown medially, yellow later-
ally, and posterior edge dark brown; legs
orangish yellow to yellow, posterior por-
tion of metatibia and entire metatarsus
yellowish brown.

Male.— As in 9 except: Head: 1.19-1.27X
as wide as long; eyes 0.63-0.71 X as wide
as long; antennae with 26-27 flagello-
meres. Mesosoma: 0.35-0.42 mm wide;
0.59-0.70 mm long; 0.57-0.65 X as wide as
long; 0.67-0.77X as wide as deep; 1.12-
1.30X as long as deep; mesoscutum usu-
ally smooth (80.0%), occasionally minutely
coriaceous (20.0%). Legs: hind tibia 0.90-
1.03 X as long as mesosoma length. Wings:
2RS 0.42-0.48 X as long as 2M. Color: me-
sosoma usually very dark brown to black
(80.0%), rarely dark reddish brown
(20.0%); tl entirely dark brown (40.0%) or
tl dark brown medially and yellow later-
ally (40.0%) or tl reddish brown medially
and yellow laterally (20.0%); t4 usually
dark brown (80.0%), rarely brown medi-
ally, yellow laterally, and posterior edge
dark brown (20.0%); remaining tergites
dark brown.

Host. — Unknown.

Material examined. — Holotype 9: USA,
South Carolina, Greenville County, Green-
ville, 18.ix.1955, G. and L. Townes (AEI).
Allotype 6 : same data as holotype except
6.vii.l952 (AEI). Other determined mate-
rial: all USA, North Carolina, Transylva-
nia County, Pisgah National Forest except
19, South Carolina, Greenville County,
Paris Mountain State Park, 16.-21. v. 1999,
R. R. Kula, Malaise trap; 19, Coontree
Creek Trail, 17.-21 .v. 1999, R. R. Kula, Mal-
aise trap; 19, Pink Beds, 17.V.1999, R. R.
Kula, sweep net; 19, 36, Coontree Creek
Trail, 18.V.1999, R. R. Kula, sweep net; 16,
Pink Beds, 19.V.1999, R. R. Kula, sweep
net (det. Kula) (TAMU).

Diagnosis. — Within the striativentris-
group O. clevelandensis Fischer, O. niobe,

296

Journal of Hymenoptera Research

and O. weemsi are morphologically similar
to O. intermissus. Head coloration can be
used to distinguish intermissus from all
three species. The head is yellow with a
very dark brown to black vertex in inter-
missus. The head is yellowish brown with
a brown vertex in clevelandensis, entirely
brown to yellowish brown in uiobc, and
mostly orange with a very dark brown
vertex and a partially brown face in weem-
si. Mesosomal coloration can also be used
to distinguish intermissus from clevelanden-
sis and niobe. In intermissus the mesosoma
is usually very dark brown to black, while
in clevelandensis and niobe the mesosoma is
brown to dark brown. Additionally, inter-
missus can be distinguished from clevelan-
densis and weemsi using exposed oviposi-
tor length and the number of antennal fla-
gellomeres, respectively. In intermissus the
ovipositor is relatively short (exposed ovi-
positor 0.18-0.32 mm long) and barely ex-
serted from the abdomen, while in cleve-
landensis the ovipositor is long (exposed
ovipositor 1.12 mm long) and conspicu-
ously exserted from the abdomen. The
number of flagellomeres in intermissus is
26-29, as opposed to 32 in weemsi.

Discussio)i. — Opius intermissus falls with-
in the striativentris-group. Dark brown me-
sosomal coloration was observed in only
one male specimen, and all other speci-
mens have a very dark brown to black me-
sosoma. Distinct color differences were
observed between female and male spec-
imens. In females t4 is dark brown medi-
ally and yellow laterally or mostly yellow
with the posterior edge dark brown. In
males t4 is usually entirely dark brown to
black. Yellow coloration on t4 was ob-
served in only one male specimen. No bi-
ological data are associated with intermis-
sus, but other species in the striativentris-
group have been reared from leaf mining
Agromyzidae (e.g. O. abbyae, O. adductus,
O. striativentris).

Opius (Gastrosema) oscinidis (Ashmead)

Rhi/ssalus oscinidis Ashmead 1889 (1888): 630.
Eutrichopsis oscinidis: Viereck 1913: 559.

Opius oscinidis: Gahan 1915: 72, 90.
Opius (Aulonotus) oscinidis: Fischer 1977: 78, 87.
Opius pusilloides Fischer 1964: 413, 433. Syno-
nym Marsh (1974).

Female. — Head: 1.21-1. 35 X as wide as
long; eyes 0.65-0.72 X as wide as long, gla-
brous; face entirely smooth to entirely ru-
gulose, slightly raised mesally, usually se-
tose (81.8%), rarely setiferous (18.2%);
clypeus flattened to convex, ventral mar-
gin lobed mesally, usually setose (81.8%),
rarely setiferous (18.2%); mandibles sim-
ple, with two apical teeth per mandible,
setiferous; antennae with 21-22 flagello-
meres. Mesosoma: 0.38-0.43 mm wide;
0.66-0.75 mm long; 0.53-0.60 X as wide as
long; 0.73-0.79 X as wide as deep; 1.34-
1.42X as long as deep; pronotum smooth
laterally, crenulate in anterior furrow; no-
tauli represented by anterior depressions;
mesoscutum smooth, setiferous anteriorly
and along lines where notauli would run
if present; mesopleuron nearly entirely
smooth, slightly to distinctly crenulate
near tegulae, sternaulus distinctly crenu-
late; scutellar disc smooth; propodeum ru-
gose, especially medially, setose; meta-
pleuron setose. Metasoma: tl 0.69-1. 00 X as
wide as long, longitudinally rugulose with
two prominent dorsal carinae delimiting a
raised median area; t2 often longitudinally
carinate at posterior edge (63.6%), occa-
sionally longitudinally carinate medially
(36.4%); t3 often smooth (63.6%), occasion-
ally longitudinally carinate at anterior
edge (36.4%); remaining tergites smooth;
exposed ovipositor 0.16-0.46 mm long.
Legs: smooth, setiferous, hind tibia 0.91-
1.00X as long as mesosoma length. Wings:
hyaline; forewing stigma narrow and
elongate; 2RS 0.42-0.59 X as long as 3RSa;
2RS 0.33-0.43 X as long as 2M; 3RSa 0.73-
0.80 X as long as 2M. Color: face and clyp-
eus brownish yellow, remainder of head
brown (54.6%) or head entirely brown
(27.3%) or face and clypeus yellow, re-
mainder of head brown (18.2%); mesoso-
ma brown to dark brown; tl dark brown;

Volume 12, Number 2, 2003

297

t2 yellow (54.6%) or yellowish brown
(45.5%); t3 often yellowish brown (63.6%),
occasionally anterior half yellow, posterior
half brown (27.3%), rarely anterior two
thirds yellow, posterior one third brown
(9.1%); t4 usually brown to dark brown
(90.9%), rarely yellowish brown (9.1%); re-
maining tergites brown to dark brown;
legs orangish yellow to yellow.

Male. — As in 9 except: Head: 1.19 X as
wide as long. Mesosoma: 0.37 mm wide.

Hosts. — An undetermined species of
CJilorops Meigen on Plantago major L. (Ash-
mead 1889); Phytomyza orobanchia Kalten-
bach (Hennig 1953); Ph. plantaginis Robi-
neau-Desvoidy (Muesebeck and Walkley
1951). All records need verification.

Material examined. — Holotype 9, O. os-
cinidis: no locality data (USNM). Holo-
type 9, O. pusilloides: USA, Allegany
County, Thomas Road near Cumberland,
28. vi. 1953, L. M. Walkley (USNM). Allo-
type 6, O. pusilloides: USA, Maryland,
Montgomery County, Cabin John,
21.viii.1917, R. M. Fouts (USNM). Para-
types O. pusilloides: all USA; 19, New York,
Otsego County, Oneonta, 24.viii.1935, H.
K. Townes; 1 9 , New York, Otsego Coun-
ty, Oneonta, 2.ix.l935, H. K. Townes; 19,
New York, Tompkins County, Ithaca,
7.ix.l935, H. K. Townes; 19, New York,
Tompkins County, Ithaca, ll.bc.1935, H. K.
Townes; 1 9 , North Carolina, Transylvania
County, Pink Beds, 22.vii.1952, G. and L.
Townes; 1 9 , North Carolina, Henderson
County, Flat Rock, 6.ix.l952, G. and L.
Townes; 1 9 , South Carolina, Greenville
County, Greenville, 24. v. 1952, G. and L.
Townes; 1 9 , South Carolina, Greenville
County, Greenville, l.ix.1952, L. and G.
Townes; 19, South Carolina, Greenville
County, Greenville, 7.ix.l952, L. and G.
Townes (AEI). Other determined material:
19, South Carolina, Greenville County,
Greenville, 1.x. 1954, G. and L. Townes
(det. Fischer as pusilloides) (AEI).

Diagnosis. — Opius oscinidis is extremely
different morphologically from all de-
scribed Nearctic species in Gastrosema. In

oscinidis t2 is at least partially longitudi-
nally carinate, although it is nearly
smooth in some individuals. Thus, oscini-
dis can be differentiated from all species in
the striativentris-group based on tergite
sculpture. Opius oscinidis lacks crenula-
tions above the episternal scrobe in the
posterior mesopleural furrow. Thus, oscin-
idis can be differentiated from all species
in the flaviceps-group based on posterior
mesopleural furrow sculpture. Addition-
ally, oscinidis has a narrowly elongate sec-
ond submarginal cell, while species in the
striativentris-group often have a narrow
but relatively short second submarginal
cell. The second submarginal cell is shape
variable for species in the flaviceps-group,
but a narrowly elongate second submar-
ginal cell, as exemplified by oscinidis, has
not been observed for species in the flavi-
ceps-group.

Discussion. — Opius oscinidis falls within
the oscinidis-group. Marsh (1974) synony-
mized O. pusilloides with oscinidis. Com-
parison of the holotypes for oscinidis and
pusilloides with a long series of determined
pusilloides from the AEI and USNM veri-
fied the synonymy. Of the 21 described
Nearctic species in Gastrosema, O. oscinidis
is the only species that falls within the os-
cinidis-group. However, several species in
other subgenera fit within the oscinidis-
group (as noted above). All biological data
associated with oscinidis needs verifica-
tion. Opius oscinidis is morphologically
similar to a species of Opius near extiratus
that has been reared from Phytomyza spp.
mining the leaves of hollies.

Opius (Gastrosema) salmonensis Fischer

Opius salmonensis Fischer 1964: 412, 438.
Opius (Gastrosema) salmonensis: Fischer 1977:
313, 385.

Female. — Head: 1. 36-1. 47 X as wide as
long; eyes 0.77-0.88 X as wide as long, gla-
brous; face punctate, slightly raised mes-
ally, setose; clypeus broadly hemiellipti-
cal, usually flattened (80.0%), rarely weak-

298 Journal of Hymenoptera Research

ly convex (20.0%), ventral margin straight posterior half dark brown (20.0%) or t3 an-
to slightly arched, often setose (60.0%), oc- terior one third orangish yellow, posterior
casionally setiferous (40.0%); mandibles two thirds dark brown (20.0%) or t3 en-
simple, with two apical teeth per mandi- tirely reddish brown (20.0%) or t3 entirely
ble, often setose (60.0%), occasionally se- dark brown (20.0%); remaining tergites
tiferous (40.0%); antennae with 21-28 fla- dark brown; legs yellow, metatibia and
gellomeres. Mesosoma: 0.61-0.86 mm wide; metatarsus usually yellow (80.0%), rarely
1.02-1.36 mm long; 0.57-0.64 X as wide as yellowish brown (20.0%).
long; 0.71-0.79 X as wide as deep; 1.25- Male. — As in 9 except: Head: face punc-
1.31 X as long as deep; pronotum smooth tate (50.0%) or rugulose (50.0%); clypeus
to rugose laterally, rugose to crenulate in flattened, setiferous (50.0%) or setose
anterior and posterior furrows; notauli (50.0%); mandibles setose; antennae with
represented by deep anterior depressions; 23-30 flagellomeres. Mesosoma: 0.56-0.67
mesoscutum often smooth (60.0%), occa- mm wide; 0.55-0.59 X as wide as long;
sionally minutely coriaceous (40.0%), usu- 0.70-0.77X as wide as deep; pronotum of-
ally setose anteriorly and along lines ten minutely coriaceous laterally (75.0%),
where notauli would run if present occasionally smooth laterally (25.0%); me-
(80.0%), rarely entirely setose (20.0%); me- soscutum often minutely coriaceous
sopleuron minutely coriaceous, deeply (75.0%), occasionally smooth (25.0%), se-
crenulate near tegulae, sternaulus broadly tose anteriorly and along lines where no-
crenulate to broadly lacunose; scutellar tauli wouM mn if present; scutenar disc
disc often smooth (60.0%), occasionally often minutel coriaceous (75.0%), occa-
minutely coriaceous (40.0%); propodeum sionall smooth (25M). metapleuron of-

areolate-rueose, setiferous; metapleuron . . Inr- no/ \ n L-c

r ,/^ on/ x 11 r ten setose (75.0%), occasionally setiferous

often setose (60.0%), occasionally setifer- nrno/\ a* 1 t1 n ™ 1 nn^, -j

,,nnnn \, , ' „ _ _. { „., (25.0%). Metasoma: tl 0.79-1.00X as wide

ous (40.0%). Metasoma: tl 0.84-1. llx as , ,cnno/, , ,L ,. n

., , n /onno/N as long, rugose (50.0%), longitudinally ru-

wide as long, usually costate (80.0%), rare- ,^_ nn,. /nr. °n, . „ J

1 nn no/ ^ ^ , , \o u 1 8ose (25.0%), or costate 25.0%); t2 costate

ly rugose (20.0%); t2 costate; t3 often Ion- ?__ __, , . ,nr-\n,^

.. .6 ,, • . • . . u ,r 4 50.0% , coriaceous (25.0% , or smooth

gitudinally carmate in anterior half and . '

smooth in posterior half (60.0%), rarely en- Wlth antenor ed§e longitudinally carinate
tirely minutely coriaceous (20.0%) or en- (25-0%); t3 smooth (50-0%)' minutelY en-
tirely smooth (20.0%); remaining tergites naceous (25.0%), or minutely coriaceous
smooth; exposed ovipositor 0.64-1.52 mm Wlth antenor edge longitudinally carinate
long. Legs: smooth, setiferous, hind tibia (250%)' remaining tergites smooth. Wings:
0.73-0.84 X as long as mesosoma length. 2RS 0-61-0.75 X as long as 3RSa; 2RS 0.43-
Wings: hyaline; forewing stigma broad 0-50x as lon§ as 2M' 3RSa 0.63-0.70 X as
and wedge shaped; 2RS 0.57-0.67X as lon§ as 2M- Color: head mostly yellow,
long as 3RSa; 2RS 0.42-0.45 X as long as vertex dark brown; mesosoma very dark
2M; 3RSa 0.67-0.75 X as long as 2M. Color, brown; tl dark brown; t2 yellow (50.0%),
head mostly yellow (80.0%) or orangish orangish yellow (25.0%), or brownish yel-
yellow (20.0%), vertex dark brown, at least l°w (25.0%); t3 anterior one third brown-
around ocelli; mesosoma usually very ish yellow, posterior two thirds dark
dark brown (80.0%), rarely reddish brown brown (25.0%) or t3 anterior half orangish
(20.0%); tl usually dark brown (80.0%), yellow, posterior half dark brown (25.0%)
rarely reddish brown (20.0%); t2 usually or t3 anterior half yellow, posterior half
yellow (40.0%) or orangish yellow (40.0%), dark brown (25.0%) or t3 anterior two
rarely reddish brown (20.0%); t3 anterior thirds yellow, posterior one third dark
half yellow, posterior half dark brown brown (25.0%); remaining tergites dark
£0.0%) or t3 anterior half orangish yellow, brown; legs yellow, metatarsus often yel-

Volume 12, Number 2, 2003

299

low (75.0%), occasionally yellowish brown
(25.0%).

Host. — Unknown.

Material examined. — Holotype 9: CAN-
ADA, British Columbia, Salmon Arm,
shore of Shuswap Lake, 13.vii.1949, H. B.
Leech (CAS). Other determined material:
26, CANADA, Province Quebec, 50°03'N
77°07'W, 12.vi.-8.viii.1987, Leblanc (AEI);
1 9 USA, New Hampshire, Grafton Coun-
ty, Mount Cardigan (USNM); 19, USA,
North Carolina, Macon County, High-
lands, 22.vi.1977, H. and M. Townes; 19,
USA, North Carolina, Macon County,
Highlands, 26.vi.1977, H. and M. Townes;
1 9 , USA, Wisconsin, Grant County, T6N
R6W SI 7, 29-v.-3.vi.1975, gypsy moth
Malaise trap; 16, USA, Wisconsin, Fond
du Lac County, T13N R19E S23, 11.-
18.vi.1975, gypsy moth Malaise trap; 16,
USA, Wisconsin, Jackson County, T21N
R4W S27, 1 6.-23. vi. 1975, gypsy moth Mal-
aise trap (det. Kula) (AEI).

Diagnosis. — Within the flaviceps-group
O. flaviceps is morphologically similar to
O. salmonensis. Opius salmonensis is con-
spicuously smaller than flaviceps. In sal-
monensis the mesosoma is 1.02-1.36 mm
long, while in flaviceps the mesosoma is
1.84 mm long. In salmonensis the mesoso-
ma is 0.56-0.86 mm wide, while in flavi-
ceps the mesosoma is 0.98 mm wide. Ad-
ditionally, exposed ovipositor length in
salmonensis is 0.64-1.52 mm, as opposed to
1.78 mm in flaviceps.

Discussion. — Opius sahnonensis falls
within the flaviceps-group. One specimen
of salmonensis in the USNM and five spec-
imens of salmonensis in the AEI were mis-
identified as O. flaviceps. All specimens
differ from the holotype of flaviceps in size
and exposed ovipositor length (as detailed
in the diagnosis for salmonensis).

Size differences were observed between
female and male specimens of salmonensis.
In females the mesosoma is 0.61-0.86 mm
wide, while in males the mesosoma is
0.56-0.67 mm wide. In females the maxi-
mum mesosoma length is 1.36 mm, and in

males the maximum mesosoma length is
1.15 mm. Differences in sculpture on t2
were also observed between the sexes. In
females t2 is always costate, while in
males t2 may be costate, coriaceous, or
smooth with the anterior edge longitudi-
nally carinate. Sculpture on t2 is generally
reduced in males relative to females.

Unfortunately, no biological data are as-
sociated with salmonensis. However, the
ovipositor of salmonensis is long relative to
species that have been reared from leaf
mining Agromyzidae (e.g. O. abln/ae, O.
adductus, O. striativentris). This suggests
that salmonensis may attack a non-leaf
miner host in a concealed situation.

Opius (Gastrosema) striativentris Gahan

Opius striativentris Gahan 1915: 72, 89.

Opius (Gastrosema) striativentris: Fischer 1977:
312, 393.

Opius gracillariae Gahan 1915: 72, 90. New syn-
onym.

Female. — Head: 1.24-1.48 X as wide as
long; eyes 0.63-0.79 X as wide as long, gla-
brous; face usually smooth with rugulose
sculpture between antennal sockets and
compound eyes (86.4%), rarely entirely
smooth (13.6%), slightly raised (60.9%) or
flattened (39.1%) mesally, setiferous; clyp-
eus hemispherical to narrowly hemiellip-
tical, convex, ventral margin straight to
slightly arched, setiferous; mandibles sim-
ple, with two apical teeth per mandible,
setiferous; antennae with 18-22 flagello-
meres. Mesosoma: 0.35-0.50 mm wide;
0.56-0.82 mm long; 0.56-0.65 X as wide as
long; 0.78-1.00 X as wide as deep; 1.31-
1.61 X as long as deep; pronotum usually
coriaceous laterally (92.1%), rarely entirely
smooth (4.6%) or dorsal half smooth ven-
tral half coriaceous (3.4%), usually crenu-
late in anterior furrow (96.6%), rarely
smooth (3.4%); notauli represented by an-
terior depressions; mesoscutum usually
smooth (94.3%), rarely rugulose medially
near transscutual articulation (5.7%), usu-
ally setiferous anteriorly and along lines
where notauli would run if present

300

Journal of Hymenoptera Research

(86.4%), rarely setiferous anteriorly only
(9.1%) or glabrous (4.6%); mesopleuron
nearly entirely smooth, often smooth near
tegulae (55.5%), occasionally rugulose to
crenulate near tegulae (44.5%), sternaulus
usually crenulate (86.4%), rarely rugulose
(10.2%) or reduced to a smooth depression
(3.41%); scutellar disc smooth; propodeum
rugulose to rugose, usually setiferous
(98.9%), rarely glabrous (1.1%); metapleu-
ron usually setiferous (98.9%), rarely gla-
brous (1.1%). Metasoma: tl 1 .00-1.56 X as
wide as long, usually rugulose (87.4%),
rarely rugose (11.5%) or smooth (1.2%); t2
coriaceous; t3 coriaceous; t4 often smooth
(59.8%), occasionally coriaceous (33.3%);
remaining tergites smooth; ovipositor
0.37-0.48 mm long. Legs: smooth, setifer-
ous, hind tibia 0.82-0.95 X as long as me-
sosoma length. Wings: hyaline; forewing
stigma narrow and elongate; 2RS 0.50-
0.69 X as long as 3RSa; 2RS 0.40-0.52 X as
long as 2M; 3RSa 0.73-0.87 X as long as
2M. Color: head entirely dark brown to or-
angish yellow with vertex dark brown to
black; mesonotum often brown to dark
brown and conspicuously darker than rest
of mesosoma (63.6%), occasionally entirely
brownish orange to dark brown (36.4%);
tl usually yellow to brownish orange
(94.3%), rarely light orangish brown
(4.6%) or yellowish brown (1.2%); t2
brownish orange to yellow; t3 brownish
orange to yellow; remaining tergites dark
brown to orangish yellow; legs yellow,
metatibia and metatarsus yellow to yel-
lowish brown.

Male— As in 9 except: Head: 1.23-1.39 X
as wide as long; eyes 0.67-0.80 X as wide
as long; face often with rugulose sculpture
between antennal sockets and compound
eyes (73.3%), occasionally entirely smooth
(26.7%). Mesosoma: 0.34-0.43 mm wide;
0.55-0.65 X as wide as long; 0.77-0.90 X as
wide as deep; pronotum usually coria-
ceous laterally (86.7%), rarely smooth
(13.3%), usually crenulate in anterior fur-
row (86.7%), rarely smooth (13.3%); me-
soscutum usually smooth (96.7%), rarely

rugulose medially near transscutual artic-
ulation (3.33%), usually setiferous anteri-
orly and along lines where notauli would
run if present (93.3%), rarely setiferous an-
teriorly only (6.7%); mesopleuron nearly
entirely smooth, often crenulate near te-
gulae (70.0%), occasionally smooth near
tegulae (30.0%), sternaulus usually crenu-
late (86.7%), rarely reduced to a smooth
depression (13.3%). Wings: 2RS 0.55-0.71 X
as long as 3RSa.

Hosts. — Phytomyza glabricola Kulp min-
ing the leaves of Ilex coriacea (Pursh) and
7. glabra (L.); P. ilicicola Loew (as P. ilicis
Curtis) mining the leaves of /. opaca So-
land. in Ait. (Langford and Cory 1937); P.
vomitoriae Kulp mining the leaves of I.
vomit or ia Soland. in Ait.

Material examined. — Holotype 9, O.
striativentris: no locality data (USNM). Ho-
lotype 9, O. graeillariae: no locality data
(USNM). Other determined material: all
USA; 1 9 , Louisiana, Cameron Parish,
Route 27, 2.1.1998, S. Scheffer; 19, Missis-
sippi, Forrest County, De Soto National
Forest, Route 13, 7.L1998, S. Scheffer; 39,
North Carolina, Montgomery County,
Uwharrie National Forest, i.1996, S. Schef-
fer; 149, North Carolina, Moore County,
Ebersole Holly Garden, 21. ii. 1996, S.
Scheffer; 6 9 , North Carolina, Wake Coun-
ty, North Carolina State University Arbo-
retum, 23.ii.1996, S. Scheffer; 19, North
Carolina, New Hanover County, Carolina
Beach State Park, 24.ii.1996, S. Scheffer;
19, North Carolina, iv.1996; 8 9, North
Carolina, New Hanover County, Carolina
Beach State Park, marina, 21.ii.1997, S.
Scheffer; 3 9, North Carolina, New Hano-
ver County, Carolina Beach State Park,
Flytrap Trail, 26.L1998, S. Scheffer; 26 9,
30 6, South Carolina, Berkeley County,
Francis Marion National Forest, Big Ocean
Bay, 18.ii.1997, S. Scheffer; 89, South Car-
olina, Charleston County, Francis Marion
National Forest, Buck Hall, 19.ii.1997, S.
Scheffer; 5 9, South Carolina, Berkeley
County, Francis Marion National Forest,
North Honey Hill Road, 19.ii.1997, S.

Volume 12, Number 2, 2003

301

Scheffer; 19, South Carolina, Berkeley
County, Francis Marion National Forest,
Road 188, 19.ii.1997, S. Scheffer; 7$, South
Carolina, Berkeley County, Francis Mari-
on National Forest, Big Ocean Bay,
21.U998, S. Scheffer; 29, Tennessee, Shel-
by County, Memphis Botanical Garden,
29.viii.1997, S. Scheffer; 19, Texas, Jasper
County, Angelina National Forest, Boykin
Springs Trail, 2.U998, S. Scheffer (det.
Kula as striativentris) (TAMU).

Diagnosis. — Within the striativentris-
group O. castaneigaster and O. hancockanus
are morphologically similar to O. striativ-
entris. Characters used to differentiate
striativentris from castaneigaster can be
found in the diagnosis for castaneigaster.
The shape of the mesosoma and tl width
to length ratios can be used to distinguish
striativentris from hancockanus. In striativ-
entris the mesosoma is 1.31-1.61 X as long
as deep and similar in shape to castanei-
gaster. In hancockanus the mesosoma is
1.26X as long as deep and similar in shape
to niobe. Tergite 1 is 1. 00-1. 56 X as wide as
long in striativentris, while tl is 0.65 X as
wide as long in hancockanus.

Discussion. — Opius striativentris falls
within the striativentris-group. In Fischer's
subgeneric classification individuals with
a crenulate sternaulus fall within Gastro-
sema, but individuals with a smooth ster-
naulus fall within Merotrachys. Opius stria-
tiventris is retained in Gastrosema because
the majority of individuals (i.e. 91.7%) fall
within Gastrosema.

Comparison of the holotype of O. gra-
cillariae with hundreds of striativentris
specimens revealed that gracillariae falls
within the morphological limits of striativ-
entris as defined in this study. Thus, O.
gracillariae Gahan 1915 is a new synonym
of O. striativentris Gahan 1915. The holo-
type of gracillariae is morphologically sim-
ilar to striativentris reared from P. glabri-
cola infesting /. coriacea and I. glabra. Opius
gracillariae and striativentris were both
originally described in Gahan (1915). Op-
ius striativentris is designated the senior

synonym because the name striativentris
has been associated with a greater number
of studies and appears more frequently in
the literature than gracillariae. Gahan
(1915) reported that gracillariae was reared
from Porphyrosela desmiodella (Clement).
This record is clearly invalid because P.
desmiodella is a lepidopteran. Opiinae, as
currently defined, is limited to braconids
that are endoparasitoids of cyclorrha-
phous Diptera. However, Porphyrosela des-
miodella feeds on Desmodium Desv., and ]a-
panagromyza desmodivora Spencer is an
agromyzid that mines the leaves of D. tor-
tuosum DC. Thus, Gahan's gracillariae may
have actually been reared from an agro-
myzid.

ACKNOWLEDGMENTS

This work was supported by National Science
Foundation Partnerships for Enhancing Expertise in
Taxonomy (PEET) grant number DEB9712543 award-
ed to R. A. Wharton and J. B. Woolley (TAMU). I am
especially grateful for the guidance and advice pro-
vided by Bob Wharton. My sincere thanks to Jim
Woolley for contributing several useful suggestions
during the course of this research. Many thanks to
Jim Ehrman (Digital Microscopy Facility, Mount Al-
lison University) for original SEMs and Greg Zolne-
rowich (Kansas State University) for advice on the
construction of plates. Special thanks to Sonja Schef-
fer (Systematic Entomology Laboratory) for provid-
ing several hundred reared specimens of O. striativ-
entris. Material supplied by the following curators
and staff members made this research possible: David
Wahl (AEI), Robert Zuparko (CAS), Henri Goulet
(CNCI), Steven Krauth (IRCW), Philip Perkins
(MCZ), Stefan Cover (MCZ), David Smith (USNM),
and Cathy Anderson (USNM).

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generic relationships within the tribe. University
of California Publications in Entomology 88: 1-112.
Wharton, R. A. 1987. Changes in nomenclature and
classification of some opiine Braconidae (Hyme-
noptera). Proceedings of the Entomological Society of
Washington 89: 61-73.
Wharton, R. A. 1988. Classification of the braconid
subfamily Opiinae (Hymenoptera). The Canadian
Entomologist 120: 333-360.
Wharton, R. A. 1997a. Subfamily Opiinae, p. 379-395.
In: Wharton, R. A., P. M. Marsh, and M. J. Shar-
key (eds.). Manual of the New World Genera of the
Family Braconidae (Hymenoptera). Special Publi-
cation of the International Society of Hymenop-
terists 1, Washington D.C.
Wharton, R. A. 1997b. Generic relationships of opiine
Braconidae (Hymenoptera) parasitic on fruit-in-
festing Tephritidae (Diptera). Contributions of the
American Entomological Institute 30: 1-53.
Wharton, R. A. 1999. A review of the Old World Ge-
nus Fopius Wharton (Hymenoptera: Braconidae:
Opiinae), with description of two new species
reared from fruit-infesting Tephritidae (Diptera).
Journal of Hymenoptera Research 8: 48-64.

J. HYM. RES.
Vol. 12(2), 2003, pp. 303-307

Prosopigastra morogoro, a New Species from Tanzania
(Hymenoptera: Apoidea: Crabronidae: Larrini)

WOJCIECH J. PULAWSKI

Department of Entomology, California Academy of Sciences, Golden Gate Park, San Francisco,
California 94118, USA; email: wpulawski@calacademy.org

Abstract. — The new species Prosopigastra morogoro Pulawski, from Tanzania, is characterized
by a densely punctate frontal protuberance and gena, an unusual female pygidial plate, presence
of a male pygidial plate, and unique male sternum VII. Its closest congener is the southern African
capensis Brauns. Several corrections are made to an earlier diagnosis of the genus by Pulawski,
1979.

I revised the world species of Prosopi-
gastra more than twenty years ago (Pu-
lawski 1979). During a collecting trip to
Tanzania in 2001, I discovered a spectac-
ular undescribed species apparently never
collected before. Its discovery requires
three corrections to my earlier diagnosis of
the genus: 1) The marginal cell is longer
than in other members of the genus and
is not broadly truncate, its length being
2.6-2.9 X maximum width of the cell (in-
ner dimensions) rather than 1.6-2.3, and
the distance between its posteroapical cor-
ner equaling 1.1-1.4X its maximum
width. Contrary to my original statement,
the cell length of the new species overlaps
with those of Holotachysphex, Parapiagetia,
and Tachysphex and therefore is not diag-
nostic for the entire genus. 2) The pygidial
plate of the female has a number of large,
ill-defined punctures on its entire surface.
The presence of an adlateral row of punc-
tures, therefore, is not diagnostic for the
entire genus. 3) Similarly, male tergum VII
has a well-defined pygidial plate and lacks
a translucent, impunctate apical depres-
sion. The presence of the depression and
lack of a pygidial plate are not character-
istics of the entire genus.

The terminology in the following de-
scription is as in Pulawski (1979).

Prosopigastra morogoro Pulawski,
new species

Name derivation. — Morogoro, a town in Tan-
zania in whose vicinity the species was first dis-
covered; a noun in apposition.

Recognition. — Prosopigastra morogoro is unique
in having a conspicuous, densely punctate fron-
tal protuberance and a uniformly, densely
punctate gena, with punctures one diameter
apart or less. In other Prosopigastra, the frontal
protuberance is either prominent and impunc-
tate or punctate and inconspicuous, and the ge-
nal punctures are several to many diameter
apart, at least near the hypostomal carina. Also,
the marginal cell of morogoro is longer than in
any other species, its anterior margin being 2.6-
2.9 X maximum cell width (inner dimensions)
rather than 1.6-2.3. The female has irregular,
large punctures on the entire pygidial plate
(Fig. Id). In the male, tergum VII has a well-
defined pygidial plate (unlike any other Proso-
pigastra), and sternum VIII is thickened near the
apex, the thickening having an apical concavity
(Fig. 2d, f), possibly a unique feature among
Apoidea.

Description. — Frons microridged between an-
tennal socket and protuberance; protuberance
prominent, punctate throughout (punctures
less than one diameter apart). Middle clypeal
section convex, with minute carina emerging
from corner of clypeal lobe and nearly parallel
to clypeal free margin; lip slightly, obtusely
pointed mesally, not incised laterally. Gena
densely, uniformly punctate throughout, punc-

304

Journal of Hymenoptera Research

-.HO

0.25 mm

••II ?'W

Fig. 1. Prosopigastra morogoro, female: a — head in frontal view, b — clypeus, c — head in lateral view, d —
pygidial plate.

tures no more than one diameter apart. Ventral
mandibular margin step-like, without preapical
expansion at distal end of notch (hence notch
open distally). Mesopleuron either punctate
(punctures less than one diameter apart) or
punctatorugose, with small unsculptured area
above scrobe. Precoxal mesopleural carina

sharp, expanded into spine in male. Propodeal
dorsum with longitudinal, anastomosed ridges,
in some specimens irregularly rugose mesally.
Marginal cell with dense microtrichia, its ante-
rior margin longer than pterostigma, with
length 2.6-2.9 X maximum cell's width (inner
dimensions), apical truncation oblique; distance

Volume 12, Number 2, 2003

305

0.25 mm

0.1 mm

0.2 mm

0.3 mm

0.1 mm

Fig. 2. Prosopigastra morogoro, male: a — clypeus, b — thorax in lateral view, c — pygidiaJ plate, d — gastral ster-
na, e — sternum VII, f — sternum VIII in oblique lateral view.

306

Journal of Hymenoptera Research

0.2 mm

Fig. 3. Prosopigastra morogoro, male genitalia: a — dorsal view, b — gonoforceps in lateral view, c — volsella, d-
penis valve.

between cell's posteroapical corner and anterior
margin equal to 1.1-1.4X cell's maximum
width. Punctures less than one diameter apart
between midocellus and orbit and on gena (ex-
cept dorsally), no more than one diameter apart
on interocellar area and scutum; less than one

diameter apart on hindfemoral outer surface
except more than one diameter apart along ven-
tral margin in female. Gastral terga coarsely
punctate, terga 1 and II each with lateral line
(lateral carina of Pulawski, 1979). Setae inclined
anterad on vertex, about 1.5 X as long as mi-

Volume 12, Number 2, 2003

307

docellar diameter, on scutum inclined posterad,
shorter than midocellar diameter (longest near
anterior margin). Mesopleural vestiture not ob-
scuring integument. Upper metapleuron large-
ly glabrous. Tergum I without tomentum in fe-
male, with rudimentary tomentum just poster-
ad of basal declivity in many males. Body black
except mandible reddish preapically in female
and yellowish white in basal two thirds in male
and tarsal apex brown. Wing membrane slight-
ly infumate, veins dark brown.

Female. — Clypeus (Fig. la): lip slightly, ob-
tusely pointed mesally, not incised laterally.
Width of postocellar area about 2.5 x length.
Precoxal mesopleural carina sharp. Pygidial
plate with irregular, large punctures, without
setae in unique specimen in unique specimen,
probably due to abrasion (Fig. Id). Length 7.5
mm.

Male. — Flagellum cylindrical. Width of pos-
tocellar area 1.7-2.1 x length. Mesothoracic ven-
ter deeply concave; precoxal mesopleural cari-
na expanded into prominent spine (which is
disproportionately larger in large specimens);
spine connected by carina to apophysis-like sig-
num. Tergum VII with well defined pygidial
plate (Fig. 2c). Sterna III— VI shallowly concave
mesally; sterna V and VI posterolaterally each
with conspicuous tuft of setae; sternum VII
with apical emargination that is partly covered
by membrane (Fig. 2e); sternum VIII conspicu-
ously thickened apically, with apical surface of
thickening characteristically concave (2f).
Length 6.1-8.5 mm. Genitalia, gonoforceps, vol-
sella, and penis valve: Fig. 3a-d.

Relationships. — Within Prosopigastra, the
lack of a ventral preapical expansion in
the mandible is shared only by morogoro
and capensis Brauns, and is clearly derived
within the genus. The markedly modified
male mesopleuron of morogoro, another
conspicuous apomorphy, is also found in
capensis and creon (Nurse). Most likely it is
an independent development in the latter
species, which belongs to the globiceps
group of Pulawski (1979). The group is

characterized by the yellow legs markings
and holoptic eyes in most males, two de-
rived features that are absent in both ca-
pensis and morogoro. Thus, capensis and
morogoro appear to be the closest relatives
within Prosopigastra.

Type material. — Holotype 6: Tanzania:
Morogoro Region: 48 km W Morogoro at
6°56.9'S 37°20.2'E, M.H. Bourbin and WJ.
Pulawski (California Academy of Scienc-
es). Paratypes (all in California Academy
of Sciences): TANZANIA: Iringa Region:
18 km W Iringa at 7°53.8'S 35°35.7'E, M.H.
Bourbin and WJ. Pulawski, 9 June 2001
(IS), 20 June 2001 (19, 56). Morogoro
Region: same data as holotype: 3 June
2001 (2 J), 6 June 2001 (2d), 11 June 2001
(26), 18 June 2001 (1 6 ); Omary S Haji and
W.J. Pulawski, 2-3 July 2001 (16), 7 July
2001 (16), and 23 July 2001 (16). Most of
the paratypes are deposited at the Califor-
nia Academy of Sciences, and one each in
the Museum fur Naturkunde, Berlin, The
Natural History Museum, London, and
the United States National Museum of
Natural History, Washington, D.C.

ACKNOWLEDGMENTS

I sincerely thank Professor Marcelian Njau (Uni-
versity of Dar es Salaam, Tanzania) for his help in
organizing my expedition, as well as my travel com-
panions in Tanzania, Mrs. Maureen H. Bourbin (Cal-
ifornia Academy of Sciences, San Francisco, Califor-
nia) and Mr. Omary S. Haji (Dar es Salaam, Tanzania)
for their help in collecting specimens; and Ms. Vir-
ginia Kirsch (San Francisco, California) for generating
the illustrations. Upon my request, Arnold S. Menke
and Michael A. Prentice reviewed earlier versions of
the manuscript and significantly improved it. Mi-
chael Ohl, the official reviewer, also contributed to
the quality of the paper.

LITERATURE CITED

Pulawski, W. J. 1979. A revision of the World Proso-
pigastra Costa (Hymenoptera, Sphecidae). Polskie
Pismo Entomologiczne 49: 3-134.

J. HYM. RES.
Vol. 12(2), 2003, pp. 308-311

A New Species of Copidosoma Ratzeburg (Hymenoptera: Encyrtidae)

from Eagle Nests in Kazakhstan

Andrey Sharkov, Todd E. Katzner, and Tatyana Bragina

(AS) Department of Biochemistry, Chemical Abstracts Service, 2540 Olentangy River Road,

Columbus, OH 43202, USA, email: asharkov2@cas.org;

(TEK) Department of Biology, Arizona State University, P.O. Box 871501,

Tempe, AZ 85287-1501, USA;

(TB) Department of Biology, Kostanay State University, 118 Taran St.,

Kostanay, 458000, Kazakhstan

Abstract. — A new species of polyembryonic encyrtid of the genus Copidosoma is described from
north-central Kazakhstan. Copidosoma naurzumense, n. sp., was reared from tineid moth larvae
collected from regurgitated pellets collected near eagle nests in the Naurzum nature reserve. This
is the third species of the genus Copidosoma reported from Kazakhstan. This species is similar to
C. longiventre Myartseva form Turkmenistan, from which it differs by having dark tegulae and
front coxae, shorter antennal segments and clava, and smaller body size. The natural history and
ecology of the parasitoid and its host are discussed.

Pe3K)Me. OnucaH hobwh bhjx 3HiiHpTH,a poaa Copidosoma H3 Ka3axcraHa

(Hayp3yMCKHH rocy^apcTBeHHbiH npHpoAHbiH 3anoBe^HHK). Copidosoma naurzumense, sp. n.
Bbme^eH H3 ryceHnu MOJieii ceMeHCTBa Tineidae (Lepidoptera), HafmeHHbix b nora/iKax,
co6paHHbix okojio rHe3^ opnoB. 3to TpeTHH npcacraBHTejib po^a Copidosoma, o6Hapy>KeHHbiH
b Ka3axcraHe. C. naurzumense sp. n. cxoneH c C longiventre Myartseva H3 TypKMeHHCTaHa, ot
KOToporo oh OTJiHHaeTca TeMHbiMH TerynaMH h nepe^HHMH Ta3HKaMH, 6ojiee KOponcHMH
HjreHHKaMH h GyjiaBoii ycHKOB, a TaioKe MeHbiiiHMH pa3MepaMH Tcna. npuBe/teHbi KpaTKne

CBeAeHHH IIO GnOJlOrHH H 3KOJIOrHH napa3HTOH,Zia H X03MHHa.

Parasitic wasps of the encyrtid genus
Copidosoma are polyembryonic parasitoids
of lepidopteran caterpillars. The genus has
a worldwide distribution and, according
to Noyes et al. (1997), includes about 150
described species. However, Trjapitzin
(1989) suggests that there are 184 de-
scribed species of Copidosoma in the world,
133 of which occur in the Palearctic. The
fauna of the family Encyrtidae of Central
Asia was revised most recently by Myart-
seva (1984), and includes 31 species of

mdosoma (4 of them as Litomastix), with
ecies, C. filicorne (Dalman) and
Ratzeburg, found in Ka-
zakhstan.

The new species described below was
reared by T.Katzner from caterpillars of
clothes moths (Lepidoptera: Tineidae) col-
lected from regurgitated pellets from ea-
gles in the Naurzumskiy Zapovednik (Naur-
zum National Nature Reserve) in the
Naurzumskiy region of the Kostanay Ob-
last' of north-central Kazakhstan (51° N,
64° E).

Copidosoma naurzumense Sharkov,
Katzner and Bragina

Female. — Body length 1.2-1.6 mm (ho-
lotype — 1.44 mm). Head: Width approxi-
mately twice its length and equal to its
height (35:18:35). Frontovertex width at

Volume 12, Number 2, 2003

309

Figs. 1-2. Copidosoma naurzumense Sharkov, Katzner and Bragina, n. sp. 1 — female antenna; 2 — male antenna.

the level of anterior ocellus approximately
Vi head width (18:35). Distance between
posterior margin of eye and occipital mar-
gin approximately 1/9 eye length from
above (1.5:13.5). Ocelli in obtuse triangle
with the angle at the anterior ocellus of
103°. POL:OOL:LOL:OCL = 11:1.3:5.5:1.
Maximum diameter of eye 1.2 times its
minimum diameter (17:14). Distance be-
tween antennal toruli twice the distance
between antennal torulus and mouth mar-
gin and approximately lA the distance be-
tween antennal torulus and eye margin (4:
2:13). Antenna as in Fig. 1. Mouth width
slightly greater than malar space (16:15).
Mesosoma: length 1.4 times its width (49:
35). Scutum transverse, its length about %
its width (22:35). Scutellum of equal
length and width (22:22). Mid tibial spur
length equal to length of first tarsomere (9:
9) and approximately V4 length of middle
tibia (9:35). Fore wing length 2.3 times its
width (93:41) (Fig. 3). Metasoma: Longer
than head and mesosoma combined. In
dry specimens its length varies depending
on degree of extension of sclerites, and

ranges from 1.1 to 2.0 times combined
length of head and mesosoma (85:60 in the
holotype). Ovipositor not exserted, with
gonostyli fused to second valvifers, their
length about Vi length of mid tibia. Color:
Head and body generally dark, almost
black. Face very dark brown, almost black,
frons and vertex black, with very slight
dark blue-green shine. Antenna dark
brown, with slightly lighter apex of the
pedicel. Scutum black, with dark blue-
green shine, which is slightly more strong-
ly expressed than on vertex; scutellum
with slight dark purple reflection. All cox-
ae dark brown. Front femur and tibia dark
brown, with very light brown apex of fe-
mur and base of tibia, tibia very slightly
lighter towards the apex; tarsus brown.
Middle femur dark brown, with light
brown apex; tibia light brown, with white
translucent base, and gradually becoming
brownish yellowish white toward apex;
spur and tarsus almost white, with last
tarsomere brownish. Hind femur dark
brown, with light apex, tibia with whitish
translucent base, dark brown in middle,

310

Journal of Hymenoptera Research

•^ •"' ^ , C ■J'^ ^

/ , /t/ • ^ ** «** ,^ U" *" '— *""^ «f^ c— v~ l — t t— tr~ t— ^ °— e— t— ^0-~ *t-

n* III / v * ■•" "^r" *■ ^ **■« _ tfc *-- ***-

Fig. 3. Copidosoma naurzumense Sharkov, Katzner and Bragina, n. sp. female forewing.

and gradually becoming brownish yellow
in apical Vr, tarsus yellowish white, with
last tarsomere brownish. Metasoma black,
with very slight metallic reflection. Sculp-
ture: Head and body reticulate, with cells
rounded on frons, vertex, and dorsal side
of mesosoma, and elongate on gena, me-
sopleuron and metasoma. Cell diameter
on dorsal side of head and mesosoma
about Vi diameter of posterior ocellus.

Male. — Body length 1.2-1.4 mm. Gen-
eral appearance as in female, except head
slightly wider (width 2.1 times length and
1.1 times height), ocelli in more obtuse tri-
angle (angle at anterior ocellus 115°), mid
tibial spur slightly shorter than first tar-
somere, and metasoma equal in length or
shorter than head and mesosoma com-
bined. Length of aedeagus, when exserted,
V3 to Vz length of mid tibia. Digiti with two
teeth. Antenna as in Fig. 2. Color and
sculpture as in female, although sculpture
patterns somewhat more pronounced.

Types.— Holotype female: 51°33.3'N

064°07.9'E KAZAKHSTAN, Kostanay re-

miskiy zapovednik, near Kar-

nerly Dokuchaevka), eagle

#10, 6.V.1998, emerged

itzner. Para types: 2 fe-

"i males, same data ex-

cept, 16.vii.1998; 30 males, same data ex-
cept, eagle pellet from nest #21, 26.V.1999;
8 females, same data except eagle pellet
from nest #13, 26.V.1999; 3 females, 4
males, same data except, eagle pellet from
nest #12, 19.vi.1999; 41 females, 43 males,
same locality, summer 1998 (deposited at
the OSU Insect Collection, Columbus,
OH).

Diagnosis. — From C. filicorne, which also
occurs in Kazakhstan, differs in having the
metasoma longer than the head and me-
sosoma combined, and dark brown front
coxa. The second species occuring in Ka-
zakhstan, C. bouchenum, has white tegulae,
the clava shorter than three preceding fu-
nicular segments, and the body length of
3.0-3.2 mm. In Myartseva's (1984) key
runs to C. longiventre Myartseva from
Turkmenistan, from which it differs by
having dark tegulae and front coxae,
shorter funicular segments and clava, and
a smaller body size. In C. longiventre fu-
nicular segments are more than twice lon-
ger than wide, clava is equal in length to
four preceding funicular segments com-
bined, and the body length is 2.4 mm. In
Trjapitzin's (1989) key C. naurzumense
runs to C. clavatutn Myartseva from Turk-
menistan, from which, according to

Volume 12, Number 2, 2003

311

Myartseva's (1984) key, it differs in having
the metasoma longer than the head and
mesosoma combined. In Kazmi and Hay-
at's (1998) key to Indian Copidosoma, runs
to C. koehleri, an introduced South Amer-
ican species, which is a parasitoid of the
potato tuber moth.

Natural history and ecology. — The climate
in the region of the Naurzumskiy Zapov-
ednik is harsh continental. Minimum win-
ter temperatures are —45 °C, and summer
maximums reach 41 °C, with average
yearly temperature being 2.4 °C. Precipi-
tation is highly variable, but averages 233
mm per year.

Parasitized and unparasitized host lar-
vae were found in summers 1997-2000 in
regurgitated pellets collected from nests
and roosts of several species of eagles Aq-
uila and Haliaeetus. Although hosts and
parasitoids were collected from pellets
during each month of the summer and
late spring, host larvae were most fre-
quently observed during the colder and
wetter months of April and May. Host lar-
vae were found in pellets composed of
mammal fur, bird feathers, or mixtures of
both. Of more than 9500 pellets evaluated,
less than 500 contained the host larvae,
with the number of larvae per pellet being
generally less than ten, but occasionally
more than 100. Current estimates of the
parasitism rate are that it is less than 40%.
Sterile defender (precocious) larvae occur
in this species, as they do in several other
polyembryonic encyrtids (Cruz 1981,
1986).

Similar numbers of tineid moths and
hymenopterans were found in pellets of
raptors in North America (Philips and
Dindal 1979). Pellets represent a large con-
centration of potential food that can sup-
port diverse invertebrate communities.

ACKNOWLEDGMENTS

The species description and drawings were made
using the facilities and equipment of the Insect Col-
lection of the Department of Entomology of the Ohio
State University. The authors are grateful to Dr. Nor-
man F. Johnson, Dr. Peter W. Kovarik, and Dr. John
W. Wenzel for their assistance. Dr. Yolanda Cruz
kindly reviewed the manuscript. Dr. John S. Noyes
(NHM) made several important suggestions. Dr. Ev-
geny Bragin, Anatoly Taran, Fergus Crystal, Seth
Layman and Doug Grier assisted with collection and
analysis of eagle pellets. This research was partially
funded by a US-EPA STAR research fellowship,
USGS Biological Resources Division, Wildlife Conser-
vation Society, Arizona State University Department
of Biology, Frank M. Chapman Memorial Fund,
Hawk Mountain-Zeiss Raptor Research Award, Ari-
zona State University Graduate Research Support
Program, The International Osprey Foundation,
World Nature Association, Arizona State University
Russian and East European Studies Consortium, and
Northwest Airlines. We acknowledge assistance of
the Naurzumskiy Zapovednik and the government of
Kazakhstan.

LITERATURE CITED

Cruz, Y. P. 1981. A sterile defender morph in a poly-
embryonic hymenopterous parasite. Nature (Lon-
don) 294: 446-447.

Cruz, Y. P. 1986. The defender role of the precocious
larvae of the polyembryonic encyrtid wasp Cop-
idosomopsis tanytmemus Caltagirone (Encyrtidae,
Hymenoptera). journal of Experimental Zoology
237: 309-318.

Kazmi, S. I. and M. Hayat. 1998. Revision of the In-
dian Copidosomatini (Hymenoptera, Chalcidoi-
dea: Encyrtidae). Oriental Insects 32: 287-362.

Myartseva, S. N. 1984. Parasitic hymenopterans of the
family Encyrtidae (Hymenoptera: Encyrtidae) of
Turkmenistan and adjacent regions of Central Asia.
Ashkhabad, Ylym, 305 p. [in Russian]

Noyes, J. S., J. B. Woolley, and G. Zolnerowich. 1997.
Encyrtidae, pp. 170-320. In: Annotated keys to the
genera of Nearctic Chalcidoidea (Hymenoptera). Ot-
tawa, NRC Research Press.

Philips, J. R. and D. L. Dindal. 1979. Decomposition
of raptor pellets, journal of Raptor Research 13:
102-111.

Trjapitzin, V. A. 1989. Parasitic Hymenoptera of the fam-
ily Encyrtidae from the Palacarctic region. Lenin-
grad, Nauka, 488 p. [in Russian]

J. HYM. RES.
Vol. 12(2), 2003, pp. 312-332

Food Plants and Life Histories of Sawflies of the Families
Tenthredinidae and Pergidae (Hymenoptera) in Costa Rica, with

Descriptions of Four New Species

David R. Smith and Daniel H. Janzen

(DRS) Systematic Entomology Laboratory, PSI, Agricultural Research Service,

U.S. Department of Agriculture, c/o National Museum of Natural History,

Smithsonian Institution, Washington, DC 20560-0168, USA, email: dsmith@sel.barc.usda.gov;

(DHJ) Department of Biology, University of Pennsylvania, Philadelphia, PA 19104, USA,

email: djanzen@sas.upenn.edu

Abstract. — Food plants and biological information are given for five species of Tenthredinidae
and six species of Pergidae reared in the Area de Conservacion Guanacaste, Guanacaste Province,
northwestern Costa Rica. The Tenthredinidae are Adiaclema chigiyae Smith, n. sp. on Entodon-
topsis leucostega (Stereophyllaceae), Waldheimia fascipennis (Norton) on Cissus pseudosicyoides (Vi-
taceae), Waldheimia suturalis (Cameron) on Cissus rhombifolia (Vitaceae), Waldheimia interstitialis
(Cameron), n. comb., on Hamelia patens (Rubiaceae), and Pristiphora auricauda Smith on Primus
annularis (Rosaceae). The Pergidae are Aulacomerus delictus Smith on Mesechites trifida (Apocyna-
ceae), Anatlntlea bimaculata (Cameron) on Hippocratea volubilis (Hippocrateaceae), Suwatnus nigriceps
(Cameron) on Psidium guajava (Myrtaceae), Acordulecera binelli Smith, n. sp., on Posoqueria latifolia
(Rubiaceae), Acordulecera liami Smith, n. sp., on Erythroxylwn havanense (Erythroxylaceae), and
Acordulecera dashielli Smith, n. sp., on Arrabidaea patellifera (Bigoniaceae). Several hundred wild-
caught larvae of these 11 species produced no parasitoids. Redescriptions are given for Aulacom-
erus daktus, Anathulea bimaculata, and Suwatnus nigriceps.

This is the second of two treatments of
the host plants and life histories of saw-
flies reared by DHJ during the Lepidop-
tera caterpillar inventory of the Area de
Conservacion Guanacaste (ACG), which
lies primarily in Guanacaste Province in
northwestern Costa Rica. The first covered
the family Argidae (Smith and Janzen
2003). Here we consider the families Ten-
thredinidae and Pergidae, the adults of
which may be distinguished in the key to
families by Smith (1988, 1995). Symphyta
larvae are keyed to family and to subfam-
ilies within the Tenthredinidae by Smith
and Middlekauff (1987). Though based on
the Nearctic fauna, this larval key will be
helpful for larvae collected in Costa Rica.
Details of the methods and rearing records
may be found at http://janzen.sas.upenn.
edu and in Janzen (2000, in press), Schauff

and Janzen (2001), Janzen et al. (2003), and
Burns and Janzen (2001).

Acronyms used are: INBio = Instituto
Nacional de Biodiversidad, Santo Domin-
go de Heredia, Costa Rica; USNM = Na-
tional Museum of Natural History, Smith-
sonian Institution, Washington, DC, USA;
BMNH = The Natural History Museum,
London, UK. Voucher numbers associated
with each reared adult are expressed as,
for example, "99-SRNP-4547"; full details
of the voucher record and associated im-
ages may be obtained at http:/ /janzen.
sas.upenn.edu.

TENTHREDINIDAE

This is a large family in the Neotropics,
with about 32 genera and over 300 species.
Four of the six Neotropical subfamilies are
known from Costa Rica, the largest being

Volume 12, Number 2, 2003

313

the Selandriinae and Blennocampinae.
Only a few species of the other two, Ne-
matinae (Smith 2003a) and Allantinae
(Smith 2003b), occur there. The subfami-
lies may be distinguished with the keys in
Smith (1995, 2003a). All Tenthredinidae in
Costa Rica have 9-segmented, filiform or
slightly compressed antennae.

SELANDRIINAE

This subfamily is recognized by its dis-
tinctive wing venation; most have vein
2A+3A complete, lack the anal crossvein,
and have vein Rs + M distinctly curved
near Sc + R in the forewing. It is the largest
subfamily of Tenthredinidae in the Neo-
tropics, and probably 60-70 species in
nine genera occur in Costa Rica. Food
plants are known for very few of them.
Most extra-tropical members of this sub-
family have ferns and grasses as food
plants.

Adiaclema Enderlein

About 30 species of Adiaclema occur
from southern Mexico to northern Argen-
tina. The genus is distinguished by the ab-
sence of the anal crossvein in the forewing
(a characteristic of the subfamily) and sim-
ple mandibles, a trait not possessed by
other Neotropical Selandriinae. All other
Neotropical selandriines have one or more
subapical teeth on one or both mandibles.
Types of all described species have been
examined by DRS except for several that
cannot be located, and it was determined
that the following species reared in Costa
Rica represents a new species. This is the
first food plant record for the genus.

Adiaclema chigiyae Smith, new species

(Figs. 1, 8, 15, 16, 27, 32, 33, 37-39)

Female. — Length, 8.0-8.5 mm. Antenna
black. Head black with clypeus, labrum,
and maxillary and labial palpi white. Tho-
rax orange. Abdomen orange with apical
three segments and sheath black. Legs or-
ange with foretarsal segments 3-4 slightly
infuscate; midfemur with a narrow black

ring at apex; midtarsus black; apical 2A of
hind tibia black with basal V3 white; hind
tarsus black. Forewing yellow with apex
beyond stigma black; costa, subcosta, stig-
ma, and veins yellow in yellow area; veins
black in black apex.

Antennal length 2.1 X head width;
length of 3rd segment 1.3X length of 4th
segment; segments beyond third gradual-
ly decreasing in length (Fig. 8). Eyes large
and converging below; lower interocular
distance 0.8 X eye length; upper interocu-
lar distance slightly greater than eye
length (Fig. 15). Head from above strongly
narrowing behind eyes; postocellar area
1.2X broader than long; distances between
hind ocelli, hind ocellus and eye, and hind
ocellus and posterior margin of head as 6:
12:10 (Fig. 16). Malar space linear; clypeus
with anterior margin slightly convex; la-
brum about 2X broader than long with
anterior margin truncate. Hind basitarsus
subequal in length to length of remaining
tarsal segments combined. Hind wing
with anal cell sessile. Tarsal claw with
long inner tooth slightly shorter than out-
er tooth and without basal lobe (Fig. 37).
Sheath short and rounded at apex in lat-
eral view; in dorsal view slightly broader
at center (Fig. 32). Lancet (Fig. 27) short,
triangular, with about 9 serrulae; protu-
berances laterally on annuli 2-10, becom-
ing more spinelike toward apex.

Male. — Length, 7.1-7.5 mm. Color simi-
lar to that of female except mesoprescu-
tum blackish at center and nearly all hind
tibia black with white only at extreme
base. Tarsal claw with long, slender outer
tooth and minute inner tooth (Fig. 38).
Genitalia (Fig. 39) with harpe elongate and
rounded at apex; parapenis narrow, taper-
ing to small rounded apex; apex of penis
valve about as long as broad, rounded at
apex, with long dorsal lobe.

Holotype. — Female, labeled "Voucher:
D. H. Janzen & W. Hallwachs, caterpillar
(Lepidoptera) database, Area de Conser-
vacion Guanacaste, Costa Rica, http://

314

Journal of Hymenoptera Research

janzen.sas.upenn.edu, 99-SRNP.9777." De-
posited at INBio.

Paratypes.— COSTA RICA: All labeled
as holotype except for voucher codes: 99-
SRNP-9757 (19); 99-SRNP-10267 (16); 99-
SRNP-10269 (19); 99-SRNP-10273 (19);
99-SRNP-10285 (1 9 ); 99-SRNP-10288 (16);
99-SRNP-10297 (1 9 ); 99-SRNP-10305 (1 9 );
99-SRNP-10314 (1 9 ); 99-SRNP-10334 (1 6 );
99-SRNP-10348 (19); Guanacaste Prov-
ince, Santa Rosa National Park, D. H. Jan-
zen, 24-VIII-14-IX-1985, Malaise trap SE-
6-C (19). Deposited at INBio and USNM.

Etymology. — This species is named in
honor of Chigiy Binell in recognition of
her great enthusiasm for the ACG and
support for the Rincon Rainforest.

Food plant and biology. — Larvae (Fig. 1)
feed on moss, Entodontopsis Icncostega
(Brid.) W. R. Buck & Ireland (Stereophyl-
laceae). In the second month of the rainy
season (late June to early July), the pen-
ultimate and last instar larvae were en-
countered feeding solitarily day and night
while fully exposed on the rough surface
of moss patches on logs and stumps in the
ACG dry forest. They were not encoun-
tered in the first two decades of the cat-
erpillar inventory because it did not occur
to us to search shady, wet patches of
moss.

The prepupal larva chews into the sur-
face of rotten bark or wood, hollows out
a smooth-walled ovoid chamber in which
to pupate, and closes the entrance hole
with wood or bark chips glued together.
There is no sign of a silken cocoon in the
chamber. The adult emerges 40-60 days
after the cocooning chamber is construct-
ed (average 47 days, n = 12) in the ACG
rainy season climate. There was no evi-
dence of pupal dormancy, but this does
not exclude the possibility that the pre-
pupae or pupae late in the rainy season
pass the six month dry season dormant in
the cocoon chamber. No parasitoids were
produced from 135 wild-caught penulti-
mate and last-instar larvae.

Remarks.— -Of the Neotropical Adiackma

species, only a few have the combination
of yellow wings with a black apex, black
head, entirely orange thorax, and orange
abdomen with the apical segments black.
This color is similar to Adiackma tetricum
(Konow) described from "Peru (Pozuzo)"
(type examined), but A. tetricum differs by
having the supraclypeal area and area
around antennal bases white, has much
longer antennae (length nearly three times
the head width), has a long, narrow ovi-
positor sheath (with valvula 3 about two
times longer than broad), has the lower in-
terocular distance about 0.9 X the eye
length, and is generally larger, being
about 10 mm in length. Adiackma blandu-
lum (Enderlein) (type not located) de-
scribed from Ecuador is also similar, but
has most of the face above the antennae
white.

In all other Adiackma species examined,
the tarsal claws of the female and male are
similar with a long inner tooth and with-
out a basal lobe (Fig. 37). It is most un-
usual for the male tarsal claws to differ so
much from those of the female (Figs. 37,
38).

BLENNOCAMPINAE

There are about 150 species in 14 genera
of Blennocampinae in the Neotropical Re-
gion and 25-30 species in five genera in
Costa Rica, most of which are in the large
genus Waldheimia. The subfamily is distin-
guished by its wing venation, with veins
M and lm-cu parallel, veins M and Rs + M
meeting Sc + R at the same point, and the
anal cell petiolate with the basal section of
vein 2A+3A absent. One species, Metape-
dias siibcoeruka (Cameron) has been reared
from Conostegia xalapensis D. Don (Melas-
tomataceae) (Smith 1995). Species of the
genus Periclista Konow are found at high
elevations in Costa Rica and may feed on
Quercus sp. (Fagaceae), as do the North
American counterparts.

Waldheimia Brulle

Nearly 100 species of Waldheimia are
known with about ten in Costa Rica, and

Volume 12, Number 2, 2003

315

the genus is widely distributed from
southwestern United States to northern
Argentina. Characteristics of the genus in-
clude the reduced apical four antennal
segments, straight forewing vein 2A + 3A,
and the bifid tarsal claws with the inner
tooth usually broader and longer than the
outer tooth and with a basal lobe. Three
species have been reared in ACG.

Waldheimia fascipennis (Norton)

(Fig. 2)

Discussion. — This is one of the few spe-
cies of Waldheimia that have the forewings
broadly black at both the base and apex
and yellow in the center. The head, thorax,
and abdomen are orange with the intero-
cellar area, apical three abdominal seg-
ments, and sheath black. The coxae, tro-
chanters, and femora are orange, the tibiae
are white at the bases with the apical half
to two-thirds black, the basitarsi are white
with a narrow black apical ring, and the
remaining tarsal segments are black. The
male is unknown.

Distribution. — Costa Rica (Guanacaste);
Mexico (Campeche, Chiapas); El Salvador.
In addition to the reared specimens, the
following from Costa Rica also have been
examined: Santa Rosa National Park, 14-
IX-5-X-1985, Malaise trap H-3-0; Prov.
Guanacaste, OTS Palo Verde Sta., 29 km
W.S.W. Canas, 10°21'N, 85°21'W, 14- VII-
1976.

Food plant and biology. — Waldheimia fas-
cipennis larvae feed on leaves of Cissus
pseudosicyoides Croat (Vitaceae) during the
rainy season in ACG dry forest (two fe-
males: 90-SRNP-1141, 83-SRNP-1144). One
record from the last month of the rainy
season represents either a second or third
rainy season generation. The bluish and
pale yellow larvae feed side-by-side on the
upper sides of the relatively horizontal
mature leaves in groups of 1-5 (Fig. 2).
They are currently indistinguishable in
color pattern, shape, and behavior to those
of Waldheimia suturalis, which have been
reared from Cissus rhombifolia in the same

habitat. Numerous Waldheimia larvae feed-
ing on both species of Cissus have not pro-
duced adults, and it may be that both spe-
cies of Waldheimia feed on both species of
Cissus.

The prepupal larva burrows down into
the litter and pupates naked in a chamber
with no silk cocoon. The adults emerged
13-14 days after the prepupa entered the
soil (n = 2). While no dormancy was re-
corded in rainy season pupae, the prepu-
pae may well pass the dry season dormant
in the litter. A total of 36 Waldheimia (W.
fascipennis and W. suturalis) wild-caught
penultimate and last-instar larvae pro-
duced no parasitoids.

Waldheimia interstitialis (Cameron),
new combination

(Fig. 3)

Discussion. — This species was described
in the genus Blennocampa Hartig by Cam-
eron (1883) and transferred to Erythraspi-
des Ashmead by Smith (in Kimsey and
Smith 1985), but Erythraspides and Wald-
heimia cannot be distinguished in the Neo-
tropics. Formal generic synonymy will be
presented later in another paper, and here
we give the new combination.

The coloration is distinctive for both
sexes of this species. The antennae are
black with the scape and pedicel orange.
The head, thorax, and abdomen are or-
ange with the interocellar area and sheath
black. The legs are orange, with the apex
of the midfemur, the entire hind femur,
the apical third to half of the mid- and
hind tibiae, and the apical three tarsal seg-
ments of all legs black. The wings are yel-
low with the apex beyond the stigma
black. Cell M is absent in the hind wing.

Distribution. — Costa Rica (Alajueia,
Guanacaste, Heredia, San Jose), Mexico
(Veracruz); Panama; Venezuela. Costa Ri-
can specimens examined are as follows:
Prov. Heredia, La Selva Biol. Sta., 3 km S
Pto. Viejo, 10°26'N, 84°01'W, 6-VI-83, 27-
IV-90, 17-1-91; Escazu, May 21, 24, 26, 27,

316

Journal of Hymenoptera Research

Figs. 1-4. Larvae. 1, Adiaclema chigiyae. 2, Waldhtimia fascipennis. 3, W. inter stitialis. 4, Aulacomerus daktus.

1987; Alajueia, 700 m, Penas Blancas, IV-
1987.

Food plant and biology. — Hamelia patens
Jacq. (Rubiaceae) was recorded as the host
plant by Kimsey and Smith (1985) who
also described and illustrated the larva
and gave the life history in Panama. One
female was reared from the same species
of food plant in Costa Rica under number
00-SRNP-9451 (three others on the same
food plant died of disease) in the lower
margin of ACG cloud forest at about 1000
m elevation (April, July). Waldheimia inter-
stitialis spun no cocoon, pupated naked in
the bottom of its rearing container, and
used eight days from prepupa until eclo-
sion. The red-headed, dark gray-blue-
black solitary larvae with the underside
yellow (Fig. 3) are quite similar to those of
the other two species of Waldheimia de-
scribed here, and, like the others, feed in

the daytime on the upper surface of the
leaf.

In laboratory rearings, Kimsey and
Smith (1985), reported that the larvae go
into the soil and form a smooth-walled,
silk-lined cocoon. There were ten days
from cocoon formation to eclosion.

Waldheimia suturalis (Cameron)

Discussion. — Both sexes of this species
are entirely black with the following parts
of the thorax red: pronotum, tegulae, me-
sonotum, mesopleuron, and upper half of
the metapleuron. The wings are uniform-
ly, darkly infuscated. The female has cell
M present in the hind wing, and the male
has a peripheral vein in the hind wing.

Distribution. — Costa Rica (Guanacaste,
Heredia); Guatemala; Honduras; Mexico
(Morelos, Veracruz). Specimens examined
from Costa Rica other than those reared

Volume 12, Number 2, 2003

317

Figs. 5-7. Larvae. 5, 6, Anathulea bimaculata. 7, Acordulecera dashielli.

318

Journal of Hymenoptera Research

8

*

13

14

Figs. 8-14. Antennae. 8, Adiaclema chigiyae. 9, Aulacomerus daktus, flagellum. 10, Anathulea bimaculata. 11,
Suwatnus nigriceps. 12, Acordulecera binelli, pedicel and flagellum. 13, A. liami, pedicel and flagellum. 14, A.
dashielli, pedicel and flagellum.

Volume 12, Number 2, 2003

319

Figs. 15-18. Head. 15, Adiacleum chigiyae, front view. 16, A. chigiyae, dorsal view. 17, Anathulea bimaculata,
front view. 18, A. bimaculata, dorsal view.

are as follows: Guanacaste Prov., OTS
Palo Verde Sta., 29 km W.S.W. Canas,
10°21'N, 85°21'W, 5-VII-1976; Heredia
Prov., La Selva Biol. Sta., 3 km S Pto. Vie-
jo, 10°26'N, 84°01'W, 17-IV-1988.

Food plant and biology. — Three females
were reared from larvae eating mature
leaves of Cissus rhombifolia Vahl (Vitaceae)
during the rainy season in ACG dry forest
(83-SRNP-727, 98-SRNP-1831, and 98-
SRNP-1833); two males were reared under
numbers 98-SRNP-l 832.1 and 98-SRNP-
1832 from the same food plant species at
the same time. The bluish and pale yellow
larvae feed side-by-side on the upper
sides of the relatively horizontal mature
leaves in groups of one to five. They are
currently indistinguishable in color pat-
tern, shape, and behavior to those of Wald-
heimia fascipennis, which have been reared
from Cissus pseudosici/oides in the same
habitat. Numerous Waldheimia larvae feed-
ing on both species of Cissus have not pro-

duced adults, and it may be that both spe-
cies of Waldheimia feed on both species of
Cissus.

The prepupal larva burrows down into
the litter and pupates in a chamber with
no silk cocoon. The adults emerged 56-102
days (average 74 days, n = 5) after the
prepupa entered the soil. This long pu-
pation period is strikingly different from
that of W. fascipennis, and probably rep-
resents some kind of rainy season dor-
mancy. As mentioned above, a total of 36
Waldheimia (W. fascipennis and W. suturalis)
wild-caught penultimate and last instar
larvae produced no parasitoids.

NEMATINAE

This is a large subfamily in the arctic,
subarctic, and temperate regions of the
Northern Hemisphere. The number of
species decreases sharply to the south, and
very few are known from the Neotropics
(Smith 2003a). Pristiphora, the only genus

320

Journal of Hymenoptera Research

known in Costa Rica, is the largest tropi-
cal-extending genus, and occurs from
Mexico south to southeastern Brazil. The
subfamily is recognized by its distinctive
forewing venation, with veins M and 1 ni-
di markedly divergent, vein M meeting
Sc + R far basal to the point where Rs + M
meets Sc + R, and the anal cell petiolate
with the base of vein 2A+3A absent.

Pristiphora Latreille

More than 50 species are known in the
Nearctic, but only nine are known from
Mexico to southern Brazil, and three in
Costa Rica, mostly from elevations above
1000 in. The Neotropical species are keyed
and described in Smith (2003a).

Pristiphora auricauda Smith

Discussion. — The distinctive coloration
of the female (antennae, thorax, and legs
black, and abdomen a contrasting bright
orange, except for black basal plates and
black anterior margin of the second seg-
ment) distinguishes this species from oth-
er Pristiphora in Costa Rica. The male has
the head, thorax, and legs mostly yellow
orange, with most of the head and thorax
dorsally and the tibiae and tarsi black. The
abdomen is orange, as is that of the fe-
male. The female ovipositor and male gen-
italia are illustrated in Smith (2003a).

Distribution. — Costa Rica (Guanacaste).
Other than the reared specimens from
ACG, an additional Costa Rican record is
from Est. Cacao, 1000-1400 m, Lado Sur-
oeste del Volcan Cacao, Prov. Guam, II
curso parataxon., Jun 1990.

Food plant and biology. — The larvae live
solitarily in leaf rolls of mature leaves of
Prunus annularis Koehne (Rosaceae). Four-
teen females and seven males were reared
from 60 leaf rolls collected from two trees
in the lower edge of ACG cloud forest at
about 1000 m elevation (00-SRNP-9074,
-9075, -9077, -9100, -9107, -9112, -9114,
-9121, -9122, -9123, -9124, -9125, -9126,
-9127, -9128, -9130, -9131, -9132, -9133,
-9134, -9135). Eleven to 30 days were used

between spinning the cocoon and eclosing
(average 17 days, n = 14) during the rel-
ative warm weather of the dry season.
There was no suggestion of dormancy.
The cocoon is a rough dark brown shaggy
cylinder spun directly on the surface of
the green leaf in the rearing container,
though in nature the larvae probably de-
scend to the litter to spin their cocoons. No
parasitoids were reared from 60 wild-
caught larvae.

PERGIDAE

Neotropical Pergidae were treated by
Smith (1990). Eight subfamilies, 32 genera,
and 256 species were recognized. Some
additional species have been described
since, and it would not be surprising if the
number of described species is eventually
doubled. Five subfamilies, ten genera, and
40 or more species occur in Costa Rica
(Smith 1995). Representatives of two sub-
families, Loboceratinae and Acorduleceri-
nae, have been reared by the ACG cater-
pillar inventory project.

Food plants for other species include jel-
ly fungus (Auricularia sp.) growing on rot-
ting wood for Decameria rufiventris (Cam-
eron) (Perreyiinae) (Smith 1995) and dried
leaves for Perreyia tropica (Norton) (Perre-
yiinae), the larvae of which travel in
groups on the ground (Flores et al. 2000).

LOBOCERATINAE

Three genera were treated by Smith
(1990), but most species are in the genus
Aulacomerus, which is the only Costa Rican
genus. Two species were recorded from
Costa Rica by Smith (1990), but more oc-
cur in the country. In Costa Rica, the sub-
family is recognized by the 7-segmented,
slightly clavate antennae with a large sen-
sory pit on the apical segment, the midtib-
iae with a preapical spine, and the hind
tibiae lacking a preapical spine.

Aulacomerus Spinola

This genus contains 22 species and oc-
curs from Mexico to northern Argentina

Volume 12, Number 2, 2003

321

Figs. 19-22. Head. 19, Suwatnus nigriceps, front view. 20, S. nigriceps, dorsal view. 21, Acordulecera binelli,
front view. 22, A. binelli, dorsal view.

(Smith 1990). This is the first food plant
record for a member of this genus.

Aulacomerus daktus Smith

(Figs. 4, 9)

Discussion. — We redescribe this species
since it was originally known from only a
few specimens. It is distinguished from
other Aulacomerus species by the key in
Smith (1990).

Description. — Length of female, 8.5-9.5
mm; male 7.0-8.0 mm. Antenna black
with scape and pedicel orange yellow.
Head orange with ocellar area and posto-
cellar area black and with light black
markings toward antenna and sometimes
from lateral ocellus to eye. Antennal tu-
bercles, supraclypeal area, clypeus, la-
brum, and mandible whitish. Thorax or-
ange. Legs orange with about apical half
of hind tibia and all hind tarsus black;
fore- and midtarsi black except bases of
basitarsi usually orange yellow. Abdomen

orange with center of 6th tergum and seg-
ments 7 to apex and sheath entirely black.
Wings yellow with apex beyond apex of
stigma black; veins and stigma yellow,
veins black in black apices.

Antennal length 1.6X head width; large
sensory area on apical segment; 3rd seg-
ment slightly longer than 4th segment
(Fig. 9). Clypeus with slight central circu-
lar emargination. Eyes slightly converging
below, lower interocular distance sube-
qual to eye length, upper interocular dis-
tance 1.1 X eye length. Head from above
narrowing behind eyes. Postocellar area
1.6X broader than long. Distances be-
tween hind ocelli, hind ocellus and eye,
and from hind ocellus to posterior margin
of head as 10:11:16. Hind basitarsus 1.3X
length of remaining tarsal segments com-
bined. Inner hind tibial spur 0.8X length
of hind basitarsus. Female lancet and
sheath and male genitalia illustrated by
Smith (1990, figs. 338, 345, 368).

322

Distribution.— Costa Rica (Guanacaste),
Guatemala, Mexico (Chiapas). This is the
first record for Costa Rica. An additional
record, other than the ACG specimens is:
Vicinity Estac Murcielago, 8 km SW Cu-
ajniquil, Guanacaste Prov., 100 m, Jun
1989, GNP Biodiversity Survey 320300,
380200.

Food plant and biology.— Adults were
reared from larvae eating mature leaves of
Mesechites trifida (Jacq.) Mull. Arg. (Apo-
cynaceae) [89-SRNP-500 (1$); 92-SRNP-
2490 (29, 16); 92-SRNP-3901 (39); 94-
SRNP-9444 (59, 16); 94-SRNP-9529 (16)].
This species is occasionally encountered in
groups of 3-7 greenish-black larvae (Fig.
4) feeding side-by-side on the upper or
lower side of a single leaf of its herbaceous
vine food plant during the mid to late
rainy season in ACG dry forest. After con-
suming the entire leaf, the group moves
up the stem to then consume the next leaf;
in larval Lepidoptera, this has been inter-
preted a a strategy to remove the visual
evidence of leaf damage from the view of
avian predators (Heinrich 1993). As is
commonplace with species of caterpillars
feeding on latex-rich plants, the larvae cut
the petiole of the leaf partly through be-
fore feeding on the blade, a behavior that
reduces the flow of fresh latex from the
bitten leaf blade (Dussourd and Eisner
1987). If the feeding group of penultimate
or last-instar larvae is molested, they walk
off in different directions on the food
plant, but within an hour they regroup
into the same feeding groups as before.

The penultimate instar larva molts into
a non-feeding orange-purple morph that,
in captivity, wanders on the foliage and
litter for about 24 hours before spinning
its smooth-walled, dark brown, ovoid co-
coon in the litter. The adult emerges 12-
16 days after cocoon spinning (average 14
days, n = 14). A total of 28 wild-caught
late instar larvae produced no parasitoids.

ACORDULECERINAE

This is a large subfamily with many un-
described species. Smith's (1990) key cov-

JOURNAL OF HYMENOPTERA RESEARCH

ers ten genera, four of which occur in Cos-
ta Rica (Smith 1995). Many are very small,
no more than 3-4 mm in length. The an-
tennae are 6-9 segmented, mostly 6-seg-
mented and filiform, the eyes are large, oc-
cupying much of the head, the mid- and
hind tibiae each have a preapical spine,
and the hind wing usually has the veins
forming the base cells RS and M in a
straight or almost straight line. Acordule-
cera, as defined by Smith (1990), is by far
the largest genus in this subfamily.

Anathulea Malaise

Anathulca is characterized by the 6-seg-
mented antenna, truncate clypeus which
is two times or more broader than long,
and the long pedicel which is one and
one-half times or more longer than broad
and nearly as long as the first flagellar seg-
ment (Fig. 10). Four species were listed by
Smith (1990) from Guatemala and Brazil,
but 10 to 20 species probably occur in the
Neotropics. This is the first food plant re-
cord for a member of this genus.

Anathulca bimaculata (Cameron)

(Figs. 5, 6, 10, 17, 18, 28, 33, 40)

Discussion. — This species has not been
treated since its original description. It
was described from Guatemala (Cameron
1883).

Female. — Length, 6.3-7.5 mm. Antenna
black with scape and base of pedicel or-
ange yellow. Head black, orange yellow
from halfway between ocelli and antennae
to clypeus and labrum. Thorax orange yel-
low with prescutum (except for sides) and
lateral lobes (except for depressed lateral
areas) black. Legs orange with apical 4 tar-
sal segments black. Abdomen orange with
segments 7 or 8 to apex black. Forewing
yellow, apex beyond apex of stigma black;
veins and stigma yellow, veins black in
black apex.

Antennal length 1.4X head width; ped-
icel 2.7 X longer than broad and 0.6 X
length of 3rd segment, 3rd segment 1.5X
length of 4th segment; long seta at apex of

Volume 12, Number 2, 2003

323

W-

JJf

3 M "

Xm y

3*

■m

p]

Pg-v A

' - * • V . ■

wtllF

. it

J

EjMU^r^SiL^

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Tf v

:JF

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23

Figs. 23-26. Head. 23, A liami, front view. 24, A. liamiA dorsal view. 25, A. dashielli, front view. 26, A dashielli,
dorsal view.

apical segment (Fig. 10). Eyes large and
strongly converging below, lower intero-
cular distance 0.6 X eye length, upper in-
terocular distance 0.9 X eye length (Fig.
17). Head from above strongly narrowing
behind eyes; postocellar area 2.1 X broader
than long; distances between hind ocelli,
hind ocellus to eye, and hind ocellus to
hind margin of head as 7:6:8 (Fig. 18).
Hind basitarsus 1.2 X longer than length of
remaining tarsal segments combined; in-
ner hind tibial spur 0.6 X length of hind
basitarsus; hind tarsal segments 2-4 each
as broad as long. Sheath (Fig. 33) long,
length subequal to length of hind tibia;
without scopae, in dorsal view broad,
rounded to acute apex. Lancet (Fig. 28)
with about 32 serrulae, broad at base and
tapering to apex, with numerous, closely
set annuli slanted toward apex dorsally.

Male. — Length, 4.8-6.0 mm. Similar to
female except mesoprescutum mostly or-
ange and costa and subcosta of forewing
more brownish. Genitalia (Fig. 40) with
harpe round, slightly broader than long,

with long hairs; parapenis acute on me-
son; penis valve rounded ventrally, slight-
ly concave dorsally, without spines.

Distribution. — Costa Rica (Guanacaste),
Guatemala, Panama. These are the first re-
cords for Costa Rica and Panama. In ad-
dition to the rearings from ACG, speci-
mens from Costa Rica have been exam-
ined from the following: Est. Maritza, 600
m, Lado oeste Volcan Orosi, Prov. Guan.,
II curso Parataxonomos, Ago 1990, L-N-
326900, 373000; Guanacaste, W side Vol-
can Orosi, Est. Maritza, 600 m, 1988;
Guanacaste NP, riparian, Oct. 20, 1977;
Guanacaste Prov., Santa Rosa National
Park, D. H. Janzen, ll-V-l-VI-85, Malaise
trap BH-9-0, 13-VII-3-VIII-85, Malaise trap
BH-12-C, 16-XI-7-XII-85, Malaise trap BH-
10-C, 24-VIII-14-IX-84, Malaise trap SE-
5-0.

Food plant and biology. — The larvae feed
on the new shoot tips and shoot epidermis
of Hippocratea volubilis L. (Hippocratea-
ceae) in the ACG dry forest (209 and 10c? :
01-SRNP-15925 to -15983, 01-SRNP-15940

324

Journal of Hymenoptera Research

to -15945, 01-SRNP-15947 to -15949, 01-
SRNP-15951 to -15953, and 01-SRNP-
16144 to -16146). The gray-brown early in-
star larvae occur in a tight cluster of 8-12
larvae arrayed on the lengthening shoot
tips (Figs. 5, 6) of vine shoots growing hor-
izontally (in search of insulated areas)
across the forest floor in the deep shade of
old-growth forest in the second month
(July) of the rainy season. In this position,
the larvae are extremely cryptic, appear-
ing to be a cluster of small leaf buds at the
shoot tip end (possibly an example of col-
lective mimicry; Pasteur 1982). The larvae
collectively eat the very tender shoot tip
back down the stem until reaching the
woody stem, and then continue back
down the stem eating off the green tender
outer epidermis, leaving the dead beige
woody branch end (Fig. 6). These later in-
star larvae, arranged 3-6 around the stem
look like dead tissue peeling back from
the stem. There is no sign of larvae in the
canopy tens of meters above where the
mature woody vines have their crowns in
the full sun.

The prepupal larvae drop or crawl off
into the litter and spin solitary beige-
brown ovoid cocoons among the dead
leaves. There are 14-20 days between co-
coon spinning and adult eclosion (average
16 days, n = 31). About 100 wild-caught
larvae of all ages produced no parasitoids.

Suwatnus Smith

This genus is very similar to Acordule-
cern, but has 7-segmented antennae (Fig.
11). One species was treated by Smith
(1990).

Suwatnus nigriceps (Cameron)

(Figs. 11, 19, 20)

Discussion. — We redescribe this species
because the original description is not ad-
equate. It was described from "Mexico"
by Cameron (1883).

Female. — Length, 4.2-4.7 mm. Antenna
black. Head black with clypeus and
mouthparts yellow orange and apical

maxillary palpal segment blackish; apex of
mandible reddish brown. Thorax orange;
tegula black; upper part of mesopleuron
may be blackish. Legs orange white with
tarsi and outer surfaces of tibiae black.
Abdomen orange ventrally and laterally,
black above with central longitudinal or-
ange stripe; sheath black. Wings lightly,
uniformly infuscated; veins and stigma
black.

Antennal length 1.1 X head width; 3rd
segment 0.8 X length of 4th segment, fla-
gellar segments each longer than broad;
apical segment without a long seta (Fig.
11). Eyes strongly converging below, lower
interocular distance 0.8 X eye length; upper
interocular distance subequal to eye length
(Fig. 19). Head from above strongly nar-
rowing behind eyes; postocellar area 2.0 X
broader than long; distances between hind
ocelli, hind ocellus and eye, and hind ocel-
lus and posterior margin of head as 11:8:19
(Fig. 20). Hind basitarsus 1.3X longer than
length of remaining tarsal segments com-
bined. Sheath with slender, posteriorly pro-
jecting scopae; sheath and lancet illustrated
by Smith (1990, figs. 415, 420).

Male. — Length, 4.3 mm. Black, with
head similar to that of female; thorax
black with pronotum brownish; legs white
with bases of coxae, tarsi, and stripe on
outer surface of mid- and hind femora
black; abdomen black with paired brown-
ish spots on terga 2-4. Genitalia illustrated
by Smith (1988, fig. 418).

Food plant and biology. — Three females
and one male were reared from larvae
feeding on mature leaves of Psidium gua-
java L. (Myrtaceae) (99-SRNP-3146) on the
ACG interface of dry forest with rainfo-
rest. The white, solitary, globular silk co-
coons were spun among the litter in the
rearing container. Six wild-caught larvae
produced no parasitoids.

Acordulecera Say

This large genus occurs from southern
Canada to Argentina. Smith (1990) did not
give a species key but listed the 45 de-

Volume 12, Number 2, 2003

325

27

9 v ^^P^^^^^^

28

Figs. 27-31. Female lancets. 27, Adiaclema chigiyae. 28, Anathulea bimaculata. 29, Acordulecera binelli. 30, A Iwmi
31, A. dashielli.

scribed species from south of the United
States. There are numerous undescribed
species, and the size of the genus may ex-
ceed 200 species. Most species are small,
4-6 mm long, and all have 6-segmented
antennae (Figs. 12-14). Probably more
than 20 species occur in Costa Rica, most
of which are undescribed. All types of the

Neotropical species have been examined
by DRS; the following three species do not
agree with the three described species
from Mexico and Central America, and
they are not the same as any of the species
described from South America. Because of
the new host plant and life historv infor-
mation, they are described here.

326

Journal of Hymenoptera Research

Acordulecera binelli Smith, new species

(Figs. 12, 21, 22, 29, 34)

Female.— Length, 5.5 mm. Antenna
black with scape orange yellow. Head
black, yellow orange below line just in
front of front ocellus (Fig. 21); mouthparts
yellow orange; apex of mandible reddish
brown. Thorax yellow orange with pro-
notum white and mesonotal front and lat-
eral lobes, tegula, and metanotum black.
Legs orange with apical three tarsal seg-
ments black. Abdomen and sheath orange.
Wings uniformly hyaline; forewing with
apical half of costa and stigma, except for
black extreme apex, yellow orange, other
veins black.

Antennal length 1.1 X head width and
2.1 X distance between eyes above; 3rd
segment 1.4X length of 4th segment; api-
cal segment slightly longer than 4th seg-
ment and 4.1 X longer than broad; apical
segment with long apical seta; hairs long,
mostly longer than width of segments
(Fig. 12). Head with scattered punctures.
Eyes converging below, lower interocular
distance about 0.7X eye length; upper in-
terocular distance about 0.9 X eye length
(Fig. 21). Head from above strongly nar-
rowing behind eyes; postocellar area
about 2.0 X broader than long; distances
between hind ocelli, hind ocellus and eye,
and hind ocellus and posterior margin of
head as 7:7:10 (Fig. 22). Distance between
antennae subequal to distance between
antenna and eye. Hind basitarsus 1.4X
length of remaining tarsal segments com-
bined. Sheath (Fig. 34) with stout, short
lateral scopae, in lateral view scopae
rounded and about equal to length of in-
ner portion of sheath, in dorsal view with
long hairs slightly curved inward. Lancet
(Fig. 29) long, with about 24 rounded ser-
rulae, serrulae at apex flatter; annuli on
apical half with hairs, annuli on basal half
with no or only short, indistinct hairs.
Male. — Unknown.

Holotype. — Female, labeled "Voucher:
D. H. Janzen & W. Hallwachs caterpillar

(Lepidoptera) database, Area de Conser-
vacion Guanacaste, Costa Rica, http://
janzen.sas.upenn.edu," "00-SRNP-9024."
Deposited at INBio.

Etymology. — This species is named in
honor of Rich Binell in recognition of his
great enthusiasm for the ACG, love of its
roads, and support for the Rincon Rain-
forest.

Food plant and biology. — Three larvae
were found feeding side-by-side on ma-
ture leaves of Posoqueria latifolia (Rudge)
Roem. & Schult. (Rubiaceae) in lower
ACG cloud forest at about 1000 m eleva-
tion. The larvae are yellowish at the rear
and thorax, with a black head. Two died
of disease and one spun a broadly ovoid,
pinkish-beige cocoon directly on the sur-
face of a leaf in the rearing container. The
adult emerged 26 days later.

Remarks. — The color and presence of a
seta on the apical antennal segment is
quite similar to Acordulecera lituratus (Ko-
now) described from Amapa, Brazil. In A.
lituratus, the abdomen is black above, and
the apical seta of the apical antennal seg-
ment is extremely long, nearly as long as
the apical antennal segment. Characters of
the antenna, head, sheath, and lancet
should be examined and compared with
the illustrations for identification of this
species.

Acordulecera liami Smith, new species

(Figs. 13, 23, 24, 30, 35, 41)

Female. — Length, 4.1-4 .4 mm. Antenna
black, scape and pedicel more whitish.
Head black with clypeus and mouthparts
yellow orange; apex of mandible reddish
brown. Thorax orange. Legs yellow or-
ange with apical 2-3 segments of fore- and
midtarsi and entire hind tarsus black. Ab-
domen orange, dorsally black with medi-
an, longitudinal orange stripe; sheath
black. Wings hyaline; veins and stigma
black.

Antennal length 0.6 X head width and
1.2 X distance between eyes from above;
3rd segment 1.4X length of 4th segment;

Volume 12, Number 2, 2003 327

apical segment subequal in length to 4th Guanacaste Province, Santa Rosa National

segment and 2.8X longer than broad at its Park, D. H. Janzen, ll-V-l-VI-1985, Mal-

greatest width; long seta at apex of apical aise trap SE-8-C (59), Malaise trap H-1-0

segment; hairs equal to or longer than (19), Malaise trap SE-5-0 (19), Malaise

width of segments (Fig. 13). Head shining trap H-2-C (1 9 ), Malaise trap SE-7-0 (3 9 ),

with scattered punctures. Eyes converging Malaise trap SE-6-C (119), Malaise trap

below, lower in terocular distance 0.8 X eye BH-9-0 (19); 1-22-VI-1985, Malaise trap

length; upper interocular distance sub- SE-6-C (19). Deposited in INBio, USNM,

equal to eye length (Fig. 23). Head from BMNH.

above strongly narrowing behind eyes; Etymology. — This species is named in

postocellar area 2.0 X broader than long; honor of Liam Binell in recognition of his

distances between hind ocelli, hind ocellus great enthusiasm for the ACG, potential as

and eye, and hind ocellus and posterior crocodile bait, and support for the Rincon

margin of head as 10:7:10 (Fig. 24). Dis- Rainforest.

tance between antennae 1.7X distance be- Food plant and biology. — The light green

tween antenna and eye. Hind basitarsus larvae feed on the new and expanding

subequal in length of length of remaining leaves of the understory shrub Erythroxy-

tarsal segments combined. Sheath (Fig. 35) him havanense Jacq. (Erythroxylaceae) in

with short, stout scopae, in lateral view the first month of the rainy season in ACG

scopa narrow and rounded, shorter than dry forest. They feed solitarily, one on

inner portion of sheath, in dorsal view each of the small ovoid leaves, and perch

with slightly incurved hairs. Lancet (Fig. -along the margin so as to appear to be part

30) with about 14 pointed serrulae; annuli of the leaf. In some years, such as 1992 and

strongly slanted apically toward dorsum 1998, they were extremely abundant, near-

and lacking hairs or armature. ly defoliating many of the food plant

Male. — Length, 3.1 mm. Similar in color shrubs. The solitary beige to brown ovoid

and structure to female except antenna cocoons are spun among the litter. Only

with only scape whitish; thorax black with five days transpire between spinning and

posterior margin of pronotum yellow or- adult eclosion. No parasitoids were reared

ange with small spot at center of meso- from hundreds of wild-caught larvae,

scutellum orange; abdomen black with Remarks. — This species is close in color

basal sterna pale orange and median Ion- to Acordulecera calceolatus (Konow) de-

gitudinal orange stripe only on anterior scribed from Oyapock, Brazil, and A. cer-

half of dorsum. Genitalia (Fig. 41) with vicatus (Konow) described from Itaituba,

harpes nearly rectangular, parapenis Brazil, all sharing the presence of a long

rounded mesally, and penis valve nearly seta on the apical antenna] segment. In A.

straight ventrally, slightly concave dorsal- calceolatus, the mesoprescutum and meso-

ly, narrowly rounded at apex, and with- scutellum are dark orange, the abdominal

out spines. dorsum is black, the apical 2-3 tarsal seg-

Holotype. — Female, labeled "Voucher: ments are black, and the sheath has more

D. H. Janzen & W. Hallwachs caterpillar slender, projecting lateral scopae. In A. cer-

(Lepidoptera) database, Area de Conseer- vicatus, the anterior half of the mesopresc-

vacion Guanacaste, Costa Rica, http:// turn, apical half of the mesoscu tell urn, and

janzen.sas.upenn.edu, 98-SRNP-1825." the metascutellum are black, the abdomen

Deposited at INBio. is orange with the apical two segments

Paratypes. — COSTA RICA: Same labels black, and the tibiae and tarsi are black,

as holotype, except numbers, 98-SRNP- Identification of A. liami should be based

1823 (19), 98-SRNP-1826 (19), 98-SRNP- on comparison of the specimens with the

1827 (19), 98-SRNP-1824 (1 6 ); Costa Rica, illustrations.

328

Journal of Hymenoptera Research

^■■M\\\\^%\

Figs. 32-36. Female sheaths, lateral and dorsal views. 32, Adiaclema chigiyae. 33, Anathulea bimaculata. 34,
Acordulecera binelli. 35, A. liami. 36, A. dashielli.

Acordulecera dashielli Smith,
new species

(Figs. 7, 14, 25, 26, 31, 36, 42)

Female. — Length, 4.8-5.2 mm. Antenna
black with scape yellow orange. Head
black with area below line just above an-
tennae and mouthparts yellow orange
(Fig. 25); apex of mandible reddish brown.
Thorax orange with tegula black; blackish
area laterally on each mesonotal lateral
lobe. Legs orange with tibiae and tarsi
black. Abdomen orange with apical two
terga and sheath black. Wings moderately

and uniformly infuscated; veins and stig-
ma black.

Antennal length subequal to head width
and 1.9X distance between eyes from
above; 3rd segment slightly shorter than
4th segment; apical segment 0.8 X length
of 4th segment and 2.5 X longer than
broad at its greatest width; apical segment
without long seta at apex; hairs equal to
or shorter than width of segments (Fig.
14). Head shining, nearly impunctate. In-
ner margin of eyes subparallel, hardly
converging below, lower and upper inter-

Volume 12, Number 2, 2003

329

ocular distances subequal to slightly
shorter than eye length (Fig. 25). Head
from above strongly narrowing behind
eyes; postocellar area 2.1 X broader than
long; distances between hind ocelli, hind
ocellus and eye, and hind ocellus and pos-
terior margin of head as 10:8:11 (Fig. 26).
Distance between antennae 2.8 X distance
between antenna and eye. Hind basitarsus
1.3X longer than length of remaining tar-
sal segments combined. Sheath (Fig. 36)
with long, narrow projecting scopae, in
lateral view slender and rounded, much
longer than inner portion of sheath; in
dorsal view, scopae forcepslike, curving
inward at apices. Lancet (Fig. 31) short,
with about 14 serrulae, with slight dorso-
apical notch at apex. Basal 8 serrulae
broad and rounded and separated by nar-
row notch; serrulae beyond 8 small and
becoming indistinct toward apex. Annuli
strongly curved, ventral half without
hairs, dorsal half with fine hairs.

Male. — Length, 3.5-4.2 mm. Similar to
female except bases of tibiae paler and
apical maxillary palpal segments blackish.
Genitalia (Fig. 42) with harpe nearly tri-
angular, lateral and apical margins round-
ed, inner margin straight; parapenis
rounded mesally; penis valve elongate,
rounded at apex, with ventral and lateral
spines.

Holotypc. — Female, labeled "Voucher:
D. H. Janzen & W. Hallwachs caterpillar
(Lepidoptera) database, Area de Conser-
vacion Guanacaste, Costa Rica, http://
janzen.sas.upenn.edu, 96-SRNP-6815."
Deposited at INBio.

Paratypes. — COSTA RICA: Specimens
labeled as above, all beginning with 96-
SRNP-: 6811 (19); 6815 (19); 6821 (16);
6822 (let); 6824 (19); 6824.1 (19); 6825
(19); 6826 (19); 6829 (19); 6948 (19);
6948.1 (19); 6948.2 (19); 6948.3 (19);
6948.4 (19); 6948.5 (19); 6948.7 (19);
6948.8 (19); 6948.10 (19); 6948.11 (19);
6948.12 (19); 6948.13 (19); 6948.14 (19);
6948.15 (19); 6948.16 (19); 6948.17 (19);
6948.18 (19); 6948.19 (19); 6948.20 (19;

6948.21 (lc5); 6948.22 (16); 6948.23 (16);
6948.24 (19); 6948.25 (19); 6948.26 (19);
6948.27 (19); 6984.6 (19); 6984.9 (19);
Guanacaste Province, Santa Rosa National
Park, D. H. Janzen, ll-V-l-VII-1985, Mal-
aise trap H-2-C (1 9 ), ll-V-l-VI-1985, Mal-
aise trap SE-8-C (16), Malaise trap H-2-C
(16), Malaise trap SE-7-0 (lc?), Malaise
trap SE-6-C (36), 1-22-VI-1988, Malaise
trap SE-8-C (1 9 ), 14-IX-5-X-1985, Malaise
trap SE-7-0 (19), 22-VI-13-VII-1985, Mal-
aise trap H-4-C (19). Deposited in INBio,
USNM, BMNH.

Etymology. — This species is named in
honor of Dashiell Binell in recognition of
his great enthusiasm for the ACG, love of
its beaches, and support for the Rincon
Rainforest.

Food pilant and biology. — The larvae feed
on the newly expanded leaves of the
wood vine Arrabidaca patellifera (Schltdl.)
Sandwith (Bigoniaceae) in the ACG dry
forest in the first month of the rainy sea-
son. The green and yellow larvae (Fig. 7)
feed on the upper side of the nearly hori-
zontal leaves. The early instars feed as a
group, but, by the penultimate instars,
they have separated to feed solitarily. The
larvae drop to the ground to spin a very
tough brown ovoid cocoon in the leaf lit-
ter. Between seven and 17 days lapsed be-
tween spinning and adult eclosion, but
some larvae remained as dormant prepu-
pae in their cocoons throughout the five
remaining months of the rainy season and
the entire six month dry season before dy-
ing, apparently because they did not get
the right cue to eclose under laboratory
conditions. This species has been found
feeding on its common food plant only at
the beginning of the rainy season, and it
may well be univoltine in nature. No par-
asitoids were produced from 50 wild-
caught larvae of all instars.

Remarks. — This species is very similar in
coloration to Acordulecera ricatus (Konow)
described from Peru, and both lack the
seta on the apical antennal segment. How-
ever, A. ricatus has a hyaline forewing

330

Journal of Hymenoptera Research

Figs. 37-42. Tarsal claws and male genitalia, ventral view of left half of genital capsule on left, lateral view
of penis valve on right. 37, Female tarsal claw of Adiaclema chigiyae. 38, Male tarsal claw of A. chigiyae. 39,
Male genitalia of A. chigiyae. 40, Male genitalia of Aimtluilca bimaculata. 41, Male genitalia of Acordulecera liami.
42, Male genitalia of A. dashielli.

Volume 12, Number 2, 2003

331

with the base and apex of the wings black,
whereas A. dashielli has uniformly lightly
blackish wings. The sheath is distinctive
for A. dashielli, but other characters need
to be compared with the illustrations for
determination.

ACKNOWLEDGMENTS

This part of the ACG caterpillar inventory has been
supported by NSF grants BSR 90-24770, DEB 93-
06296, DEB-94-00829, DEB-97-05072, and DEB-
0072730, by taxonomists of the Smithsonian Institu-
tion and the Systematic Entomology Laboratory of
the U. S. Department of Agriculture, and by financial,
administrative, and logistic support from Costa Rica's
INBio, the government of Costa Rica, the Area de
Conservacion Guanacaste, and CONICYT of Costa
Rica. Many individuals have supported the devel-
opment of all stages of the project in a multitude of
ways. We specially thank the following people for
caterpillar hunting and husbandry: W. Hallwachs, R.
Moraga, G. Sihezar, G. Pereira, L. Rios, M. Pereira, O.
Espinosa, E. Cantillano, M. Pereira, R. Franco, H. Ra-
mirez, F. Chavarria, M. M. Chavarria, E.Olson, C.
Moraga, P. Rios, C. Cano, D. Garcia, F. Quesada, E.
Araya, E. Guadamuz, R. Espinosa, R. Blanco, A. Gua-
damuz, D. Perez, R. Blanco, F. Chavarria, C. Camar-
go, H. Kidono, A. Masis, and W. Haber.

We thank the following for allowing examination
of Costa Rican collections: J. Ugalde Gomez, Instituto
Nacional de Biodiversidad, Santo Domingo de He-
redia, Costa Rica; I. Gauld, The Natural History Mu-
seum, London, U.K.; P. Hanson, Universidad de Cos-
ta Rica, San Jose; J. Longino, The Evergreen College,
Olympia, Washington, U.S.A.; and H. Hespenheide,
University of California, Los Angeles, California,
U.S.A. Cathy Apgar, Systematic Entomology Labo-
ratory, U. S. Department of Agriculture, took the pho-
tographs of the ovipositors, heads, and antennae, and
arranged and labeled the plates. We appreciate the
reviews by N. Schiff, U.S. Forest Service, Stoneville,
MS, and S. Scheffer and E. E. Grissell, Systematic En-
tomology Laboratory, U. S. Department of Agricul-
ture, Beltsville, MD, and Washington, DC, respective-
ly-

LITERATURE CITED

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pyrrhopygine skipper butterflies (Hesperidae) in
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43.

Cameron, P. 1883. Hymenoptera, Tenthredinidae—
Chrysididae. In Godman and Salvin, Biologia
Centrali-Americana, Vol. 1, 486 pp.

Dussourd, D. E. and T. Eisner. 1987. Vein-cutting be-

havior: Insect counterploy to the latex defense of
plants. Science 237: 898-901.

Flores, C, J. Ugalde, P. Hanson, and I. Gauld. 2000.
The biology of perreyiine sawflies (Hymenop-
tera: Pergidae) of the Perreyia genus-group, pp.
258-266. In Austin, A. D„ and M. Dowton, eds.
Hymenoptera Evolution, Biodiversity and Biological
Control. CS1RO Publishing, Collingwood, Austra-
lia. 468 pp.

Heinrich, B. 1993. How avian predators constrain cat-
erpillar foraging, pp. 224-247. In Stamp, N. E.
and T. M. Casey, eds. Caterpillars, Ecological and
Evolutionary Constraints on Foraging. Chapman &
Hall, New York. 587 pp.

Janzen, D. H. 2000. Costa Rica's Area de Conserva-
cion Guanacaste: a long march to survival
through non-damaging biodevelopment. Biodi-
versity 1: 7-20.

Janzen, D. H. In press. Ecology of dry forest wildland
insects in the Area de Conservacion Guanacaste,
northwestern Costa Rica, pp. 1-44. In Frankie, G.
W., A. Mata, and S. B. Vinson, eds. Biodiversity
Conservation in Costa Rica: Learning the Lessons in
Seasonal Dry Forest. University of California
Press, Berkeley.

Janzen, D. H., A. K. Walker, J. B. Whitfield, G. Del-
vare, and I. D. Gauld. 2003. Host-specificity and
hyperparasitoids of three new Costa Rican spe-
cies of Microplitis Foerster (Hymenoptera: Bra-
conidae: Microgastrinae), parasitoids of sphingid
caterpillars. Journal of Hymenoptera Research 12:
42-76.

Kimsey, L. S. and D. R. Smith. 1985. Two new species,
larval descriptions and life history notes of some
Panamanian sawflies (Hymenoptera: Argidae,
Tenthredinidae). Proceedings of the Entomological
Society of Washington 87: 191-201.
Pasteur, G. 1982. A classificatory review of mimicry-
systems. Annual Review of Ecology and Systematica
13: 169-199.
Schauff, M. E. and D. H. Janzen. 2001. Taxonomy and
ecology of Costa Rican Euplectrus (Hymenoptera:
Eulophidae), parasitoids of caterpillars (Lepidop-
ter a). Journal of Hymenoptera Research 10: 181-230.
Smith, D. R. 1988. A synopsis of the sawflies (Hy-
menoptera: Symphyta) of America south of the
United States: introduction, Xyelidae, Pamphili-
idae, Cimbicidae, Diprionidae, Xiphydriidae, Sir-
icidae, Orussidae, Cephidae. Systematic Entomol-
ogy 13: 205-261.
Smith, D. R. 1990. A synopsis of the sawflies (Hy-
menoptera, Symphyta) of America south of the
United States: Pergidae. Revista Brasileira Ento-
mologia 34 ( 1 ): 7-200.
Smith, D. R. 1995. 6, The sawflies ^nd woodwasps,
pp. 157-177. In Hanson, P. F. and I. D. Gauld,
eds. The Hymenoptera of Costa Rica. Oxford Uni-
versity Press, Oxford, U.K. 893 pp.

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Smith, D. R. 2003a. A synopsis of the sawflies (Hy-
menoptera: Symphyta) of America south of the
United States: Tenthredinidae (Nematinae, Het-
erarthrinae, Tenthredininae). Transactions of the
American Entomological Society 129: 1-45.

Smith, D. R. 2003b. A synopsis of the sawflies (Hy-
menoptera: Symphyta) of America south of the
United States: Tenthredinidae (Allantinae). Jour-
nal of Hymenoptera Research 12: 148-192.

Smith, D. R. and D. H. Janzen. 2003. Food plants and
life histories of sawflies of the family Argidae
(Hymenoptera) in Costa Rica, with descriptions
of two new species. Journal of Hymenoptera Re-
search 12: 193-208.

Smith, D. R. and W. W. Middlekauff. 1987. Suborder
Symphyta, pp. 618-649. /;; Stehr, F. W., ed. Im-
mature Insects. Kendall Hunt Publishing Com-
pany, Dubuque, Iowa. 754 pp.

J. HYM. RES.
Vol. 12(2), 2003, pp. 333-345

Review of the Southeastern Asian Sawfly Genus Eusunoxa Enslin

(Hymenoptera: Tenthredinidae)

David R. Smith and M. S. Saini

(DRS) David R. Smith, Systematic Entomology Laboratory, PSI, Agricultural Research Service,

U.S. Department of Agriculture, % National Museum of Natural History,

Smithsonian Institution, Washington, DC 20560-0168, USA, email: dsmith@sel.barc.usda.gov;

(MSS) Department of Zoology, Punjabi University, Patiala, 147002 India

email: saini20@glide.net. in

Abstract. — Nine species of Eusunoxa Enslin are keyed, described, and illustrated: E. buchi Togashi
from the Philippines; E. ceylonica Malaise from Sri Lanka and southern India; E. formosana Enslin
from Taiwan; E. ebena, n. sp., from Indonesia (Kalimantan); and E. nigriceps (Rohwer), E. auri-
cauda, n. sp., E. lissofrons, n. sp., E. punctata, n. sp., and E. semipunctata, n. sp., from southern
India. Eusunoxa subg. Asunoxa Wei is a new synonym of Eusunoxa Enslin, and Eusunoxa In-
diana Haris is a new synonym of E. ceylonica Malaise.

Eusunoxa Enslin, a small genus of the
subfamily Allantinae, is known from In-
dia, Sri Lanka, Taiwan, Indonesia, and the
Philippines. Malaise (1963) also men-
tioned Burma in the distribution of the ge-
nus, but we have not seen specimens from
Burma, although one species was collected
during extensive surveys of northeastern
India by the junior author. Specimens are
not common, but significant collections
from southern India and Sri Lanka have
prompted this review. We treat nine spe-
cies, five of which are described as new.
Food plants are not known.

Acronyms used are: BMNH = The Nat-
ural History Museum, London, UK; DEI
= Deutsches Entomologisches Institut,
Eberswalde, Germany; PUNJ = Punjabi
University, Patiala, India; USNM = Na-
tional Museum of Natural History, Smith-
sonian Institution, Washington, DC, USA.
Abbreviations used are: OOL = distance
between eye and hind ocellus; POL == dis-
tance between hind ocelli; OCL = distance
between hind ocellus and posterior mar-
gin of head.

Eusunoxa Enslin

Eusunoxa Enslin 1911: 99. Type species: Eusunoxa
formosana Enslin, by original designation.

Eusunoxa subg. Asunoxa Wei 1997: 88. Type spe-
cies: Eusunoxa ceylonica Malaise, by original
designation. New synonymy.

Description. — Antenna filiform, some-
times slightly incrassinate in middle and
flagellomeres slightly serrate, length 2X
or less head width; 1st and 2nd segments
each longer than broad; 3rd segment sub-
equal to or slightly longer than 4th; apical
segments without ventral membranous
areas. Head from above strongly narrow-
ing behind eyes; postgenal carina absent;
inner margins of eyes slightly converging
downwards; clypeus subtruncate to very
shallowly circularly emarginate (Fig. 2);
labrum short, about 2x broader than
long; malar space linear or very narrow
to slightly more than half diameter of
front ocellus. Head with frontal area
about same height as eyes; supraantennal
tubercles and frontal ridges indistinct;
median fovea a shallow groove; supraan-
tennal pits distinct, about 2x their di-

334

Journal of Hymenoptera Research

Figs. 1-4. Front view of head. 1, Eusunoxa punctata. 2, E. semipunctata. 3, Eusunoxa ceylonica. 4, E. lissofrons.

ameter from antennal socket and each
with a small conical projection at center;
lateral furrows deep and slightly diverg-
ing posteriorly; post-, inter-, and circu-
mocellar furrows indistinct to absent. Ep-
icnemium present as a flat sclerite sepa-
rated from mesepisternum by a suture.
Hind coxae lengthened, end of hind bas-
itarsus reaching to and beyond apex of
abdomen; hind basitarsus (Figs. 5-8) lon-
ger than length of remaining tarsal seg-
ments combined, broad and laterally flat-
tened, with outer surface concave; tarsal
pulvilli present only on segments 3 and
4. Tarsal claws with single inner tooth,
slightly shorter than outer tooth, and po-
sitioned lateral to outer tooth; basal lobe
present or absent. Hind wing with cell Rs
absent, cell M present. Thorax and ab-
domen impunctate, shining.

Discussion. — The presence of an epicne-

mium, long hind legs with the hind basi-
tarsus reaching to or beyond the apex of
the abdomen, the large, laterally com-
pressed hind basitarsus, and presence of
one closed cell in the hind wing are dis-
tinctive for Eusunoxa. Eusunoxa may be
separated from other Allantinae by Mal-
aise's (1963) key. It is closest to Megabeleses
Takeuchi, Beleses Cameron, and Nesotax-
onus Rohwer, but Megabeleses has the inner
tooth of the tarsal claws much longer than
the apical one, much longer antennae
(more than twice the head width), and the
head from above slightly dilated behind
the eyes. The other genera have a normal
(cylindrical) hind basitarsus and have ven-
tral membranous areas in the apical four
antennal segments. The broad, laterally
compressed hind basitarsus is reminiscent
of the same in the Holarctic genus Craesus
Leach (Nematinae).

Volume 12, Number 2, 2003

335

Wei (1997) proposed the subgenus Asu-
noxa for those species lacking a basal lobe
to the tarsal claws (Fig. 22). This included
all known species except E. fonnosana En-
slin, which has a basal lobe (Fig. 21) and
was the only described species in the typ-
ical subgenus (Wei [1997] separated some
unnamed species, designated as "£. spp.,"
and we add one, E. ebena, n. sp., here).

Other than the basal lobe, £. formosana
shares all other characters with other spe-
cies, including similarities in the genitalia,
and shares the presence of punctures on
the frons with some other species. Because
the presence or absence of a basal lobe is
of suspicious importance, we do not see
the necessity of recognizing subgenera at
present.

KEY TO SPECIES OF EUSUNOXA

1 Tarsal claws with basal lobe (Fig. 21); front of head with large, closely set punctures

separated by narrow ridges, as in Fig. 1 2

- Tarsal claws without basal lobe (Fig. 22); punctures on front of head as above, widely

spaced with broad, flat, shining interspaces (Fig. 2) or absent (Figs. 3, 4) 3

2 Head, thorax, and abdomen orange yellow; antenna black with basal segments orange;

wings yellow hyaline, costa and subcosta yellowish, stigma brownish with margins
yellowish; hind basitarsus 4.8 X longer than maximum width (lancet in Fig. 9; male
genitalia in Figs. 13, 14) fonnosana Enslin

- Head (except for reddish-brown clypeus), thorax and abdomen black; wings lightly black-

ish with costa, subcosta and stigma black; hind basitarsus 3.8x longer than maximum
width ebena, n. sp.

3 Thorax black or black with pronotum, tegula, and part of meso- and metanotum dark

orange 4

- Thorax entirely orange

4 Pronotum, tegula, V-shaped mark on mesoprescutum, mesoscutellum, and metanotum

orange (head shining, impunctate, as in Fig. 4; male genitalia with penis valve curved,

in Fig. 20) nigriceps (Rohwer) (6)

- Thorax entirely black 5

5 Abdomen orange except for blackish basal plates (frons sparsely punctured, with shining

interspaces, as in Fig. 2; male genitalia in Fig. 17) auricauda, n. sp.

- Abdomen black 6

6 Legs entirely black; head shining, impunctate (Fig. 4); penis valve oval (Fig. 19)

lissofrons, n. sp.

- Legs with fore- and midtibiae and tarsi white; head with sparse, widely separated punc-

tures with shining interspaces on frons (Fig. 2); penis valve elongate, constricted at
center (Fig. 18) semipunctata, n. sp. (6)

7 Clypeus and supraclypeal area dark orange; first antennal segment or basal 2 antennal

segments and base of 3rd segment may be orange; head with Frons densely punctate,
punctures close together, separated bv narrow ridges, without shining interspaces (Fig.

1) 8

- Head and antenna black; frons impunctate, shining, or with widely spaced punctures

separated by broad, shining interspaces (Figs. 2—4) 9

8 Hind femur basally orange, apical half or more black; basal 2 antennal segments and base

of 3rd segment orange; hind basitarsus about 1.4X length of remaining tarsal segments
combined; Philippines buchi Togashi

- Hind femur entirely black; only 1st antennal segment dark orange; hind basitarsus about

336

Journal of Hymenoptera Research

1.7X length of remaining tarsal segments combined (Fig. 8) (lancet in Fig. 11); India

punctata, n. sp.

9 Femora blackish, only extreme bases may be orange; frons with distinct, widely spaced

punctures, separated by broad, shiny interspaces (Fig. 2) (lancet in Fig. 12)

semipunctata, n. sp. (?)

- Fore- and midfemora orange, hind femur entirely or partly black; frons with few minute

punctures or impunctate, shining (Figs. 3, 4) 10

10 Most of hind femur black, only extreme base orange; extreme apices of fore- and midtibiae

black (lancet in Fig. 10; male genitalia in Figs. 15, 16) ceylonica Malaise

- Apical half of hind femur black; tibiae orange (after Rohwer 1915) . . . nigriceps (Rohwer) (9)

Ensiinoxa auricauda Smith and Saini,
new species

(Fig. 17)

Female. — Unknown .

Male. — Length, 6.5 mm. Antenna, head,
and thorax black, abdomen orange with
black basal plates, basal plates darkest
black laterally. Legs with apices of coxae
and trochanters on upper surfaces white;
fore- and midlegs with apices of femora
and tibiae and tarsi entirely white; hind
tibia black with white streak on inner sur-
face of basal half. Wings lightly infumated
towards apex; stigma, costa, and rest of
venation dark brown to black. Antenna
slightly compressed and flagellomeres ser-
rate on underside with blunt projections
on segments at apex below; length 2X
head width; pedicel 2x its apical width;
scape as long as its apical width; segment
3 slightly longer than segment 4. Clypeus
with shallow V-shaped emargination;
lower interocular distance 0.8X eye
length; OOL:POL:OCL 1.00:0.92:0.80;

postocellar area subconvex, 3X broader
than long at its maximum width; head
with sparse, shallow punctures with flat
shining interspaces on and around the
frontal area (similar to Fig. 2), posterior or-
bits and postocellar area impunctate. Hind
basitarsus 1.3X length of remaining tarsal
segments combined, about 4.0 X longer
than maximum width. Tarsal claws with-
out basal lobe. Genital capsule similar to
Fig. 15; penis valve in Fig. 17.

Holotype. — 6, India, Kerala: Munar,

4000 ft., 22.9.1995, collection M.S. Saini
(PUNJ).

Distribution. — India (Kerala).

Etymology. — The species name is based
on the orange abdomen.

Remarks. — The black head and thorax
with the contrasting orange abdomen are
unlike other species of Eusunoxa. This col-
oration and the sparsely punctured head
(similar to Fig. 2) will distinguish this spe-
cies from other species in the genus. Eu-
sunoxa auricauda is most similar to the
male of E. semipunctata, with which it
shares the punctate head, similar hind
basitarsus, and very similar male genitalia
(Figs. 17, 18). However, the absence of col-
or variation in the large series of males of
E. semipunctata and the slight difference in
shape of the penis valve (less constricted
at its center than that of E. semipunctata),
provide evidence that E. auricauda is a sep-
arate species.

Eusunoxa buchi Togashi

Eusunoxa buchi Togashi 1981: 419, figs. 21-24,
26-28.— Wei 1997: 89 (in subg. Asunoxa; in
key).

Female. — Unknown.

Male. — Length, 6.0 mm. Antenna black
with segments 1, 2, and basal half of 3rd
segment orange. Head black with clypeus,
mouthparts, and supraclypeal area or-
ange. Thorax and abdomen orange. Fore-
and midlegs orange with apical 4 tarsal
segments infuscate; hind leg with coxa
and trochanter orange, femur black except

Volume 12, Number 2, 2003

337

8

Figs. 5-8. Apex of hind tibia and hind tarsus. 5, Eusunoxa ceylonica. 6, E. lissofrons. 7, E. semipunctata. 8, E.

punctata.

basal quarter orange, tibia black with bas-
al third white, and tarsus black. Wings
very lightly, uniformly infuscated; veins
and stigma black. Antennal length 1.7X
head width; 3rd segment subequal in
length to 4th. Lower interocular distance
0.7X eye length. Postocellar area 2.5 X
broader than long; clypeus subtruncate;
malar space linear; OOL:POL:OCL == 1.00:
0.80:0.80. Front of head densely punctate,
punctures large and close together, sepa-
rated by narrow ridges (similar to Fig. 1);
postocellar and postocular areas nearly

impunctate, shining. Hind basitarsus 1.4X
length of remaining tarsal segments com-
bined, 4.2X longer than maximum width.
Tarsal claws without basal lobe.

Types.— Holotype 6, "Sept. 7, 1961,
Mantalingajan, Pinigisan, 600 m, Palawan
Is., Noona Dan Expedition," in Universi-
tets Zoologiske Museum, Copenhagen,
Denmark. Ten paratype males, dated Sept.
6-24, 1961, also in type series. None ex-
amined.

Specimen examined.— PHILIPPINES: Pa-
lawan, Mantalingajan, Pinigisan, 600 m, 23

338

Journal of Hymenoptera Research

Sept. 1961, Noona Dan Exp. 61-62 (1 6\
BMNH). From same locality as type series.

Distribution. — Philippines (Palawan).

Remarks. — Eusunoxa buchi is the only
species in the genus known from the Phil-
ippines. It shares the densely punctate
head with E. formosana, E. ebena, and £.
punctata. The mostly black coloration and
lack of a basal lobe on the tarsal claws sep-
arate it from E. formosana and E. ebena, and
the dark orange basal antennal segments
and only the apical half of the hind femur
black separate it from E. punctata from In-
dia, which has the hind femur black and
only the first antennal segment dark or-
ange.

Eusunoxa ceylonica Malaise

(Figs. 3, 5/l0, 15, 16, 22)

Netrocerus nigriceps Enderlein 1920: 371. Preoc-
cupied in Eusunoxa by Eusunoxa nigriceps
(Rohwer 1915).

Eusunoxa ceylonica Malaise 1932: 147. — Smith
1982: 120,' figs. 2, 8, 14 (Sri Lankan records;
female lancet, male genitalia, tarsal claw fig-
ured; syn.: N. nigriceps Enderlein). — Saini
and Deep 1994: 50 (India, Tamil Nadu; as
"cevlonica").—Wei 1997: 89 (in subg. Asu-
noxa; in key).

Eusunoxa indiana Haris 2000: 299. New synon-
ymy.

Female. — Length, 6.5-7.5 mm. Antenna
and head black; clypeus black to partially
orange on side or anterior margins. Tho-
rax and abdomen orange. Foreleg orange
with extreme apex of tibia and apical 4
tarsal segments blackish; midleg orange
with apical half of tibia and entire tarsus
black; hind leg with coxa, trochanters, and
basal third of femur orange, apical two-
thirds of femur and entire tibia and tarsus
black. Wings uniformly infuscated; veins
and stigma black. Antennal length 1.7X
head width, 3rd segment slightly longer
than 4th. Lower interocular distance sub-
equal to eye length; clypeus subtruncate;
postocellar area 4.5 X broader than long;
malar space nearly linear; OOL:POL:OCL
= 1.00:0.90:0.35. Head shining, impunctate

except for small widely scattered punc-
tures on frontal area (Fig. 3). Hind basi-
tarsus (Fig. 5) 1.5X length of remaining
tarsal segments combined; about 3.6 X lon-
ger than maximum width. Tarsal claws
without basal lobe. Lancet in Fig. 10, with
about 17 serrulae; apex roundly acute.

Male. — Length, 5.5-6.0 mm. Similar to
female but with terga 2 to apex mostly in-
fuscate to blackish and clypeus black or
partly to almost entirely orange. Genitalia
in Figs. 15, 16.

Types. — The holotype 9 of Netrocerus ni-
griceps Enderlein, is in the Polish Academy
of Sciences, Warsaw (examined). The ho-
lotype 9 of £. ceylonica Malaise is in the
BMNH, labeled "Colombo, Ceylon"; a
paratype from the Swedish Museum of
Natural History was examined. The holo-
type of £. indiana Haris is at the BMNH
(examined), a 6 labeled: "Holotype," "In-
dia, Tamil N., Madras, 2.XI.79. Boucek,"
"Holotype Eusunoxa indiana sp. n. det. A.
Haris 1999," "B.M. type Hym. 1.851."

Specimens examined. — INDIA: Poonmu-
di, 18-9-95 (1 9); South India, Anamalai
Hills, Cinchona, 3500 ft., V-1964, P.S. Na-
than (2 9), V-1966 (2 9), X-1966 (1 9), V-
1967 (1 9), V-1968 (1 9); Nilgiri Hills, Sin-
gara, S. India, 3400 ft., June 1954, col: P.S.
Nathan (1 9); South India, Kerala State,
Trivandrum Dt., Poonmudi Range, May
1971, 3000 ft. elevation, T.R.S. Nathan (2
9); South India, Pondicherry State, Kari-
kal, P.S. Nathan (1 9 ), same but with date,
X-'62 (1 9). SRI LANKA: Western Prov-
ince, Central Province, Southern Province,
Eastern Province, North Western Prov-
ince, Northern Province (see Smith 1982
for records); Ceylon, Trincomalee (1 9,
BMNH).

Distribution. — India (Kerala; Tamil
Nadu); Sri Lanka.

Remarks. — Eusunoxa indiana Haris is the
male of £. ceylonica. It is typical except for
the orange clypeus. We have not seen oth-
er specimens with an entirely orange clyp-
eus; however, among the material we ex-
amined, coloration varies from black to

Volume 12, Number 2, 2003

339

%^^ l-

f^

■I '

12

Figs. 9-12. Female lancets. 9, Eusunoxa formosana. 10, E. ceylonica. 11, E. punctata. 12, E. semipunctata.

brownish, to some partially orange at the imen and glued on cardboard on the same

base and laterally. pin) is more punctate than most spuci-

The specimen from Ceylon at BMNH is mens,

like E. ceylonica, but the head (off the spec- We have not seen variation in color of

340

Journal of Hymenoptera Research

the thorax and abdomen, both are mostly
orange, with the abdomen of the male
more blackish dorsally. Eusunoxa ceylonica
appears to be most similar to E. nigriceps,
both sharing the shining head with few
widely spaced punctures, and the female
is difficult to distinguish from that of E.
nigriceps based on Rohwer's (1915) de-
scription of the latter. The male paratype
of E. nigriceps, however, is mostly black,
and the genitalia differ (Figs. 16, 20) from
that of E. ceylonica.

In Sri Lanka, this species occurs in both
open and wooded areas in dry and wet
zones with annual rainfall ranging from
660 to 1952 mm (Smith 1982).

Eusunoxa ebena Smith and Saini,
new species

Female. — Length, 8.0 mm. Antenna
black with segments 1, 2, and basal half of
3 dark orange; head black with clypeus
reddish brown. Thorax and abdomen
black. Foreleg dark brown with coxa and
apical 3 tarsal segments black; midleg
with coxa black, femur and tibia dark
brown (tarsus missing); hind leg black
with basal half of tibia white (apical two
tarsal segments missing). Wings very
lightly, uniformly infuscated; veins and
stigma black. Antennal length 1.6x head
width, 3rd segment 1.1 X longer than 4th.
Lower interocular distance 0.8 X eye
length; clypeus very shallowly circularly
emarginated; postocellar area 2.7X broad-
er than long; malar space distinct, slightly
more than half diameter of front ocellus;
OOL:POL:OCL = 1.0:1.2:1.0. Head with
frontal area densely punctured, punctures
deep, close together, separated by narrow
ridges (similar to Fig. 1); clypeus impunc-
tate, shining; postocellar area and head be-
hind eyes sparsely punctate, shining. Hind
basitarsus extremely flat and concave;
3.8 X longer than maximum width (length
to remaining tarsal segments unknown
since apical tarsal segments missing). Tar-
sal claws with basal lobe (Fig. 21).

Male. — Unknown.

Holotype. — $, "Borneo, Pontianak, F.
Muir" (BMNH).

Distribution. — In present-day Kaliman-
tan, Indonesia.

Etymology. — The name is based on the
almost entirely black coloration of this
species.

Remarks. — Eusunoxa ebena and E. formo-
sana are the only species that have a basal
lobe on the tarsal claws. Also, both share
the densely punctate frons. The coloration
of the two are, however, entirely different.
Eusunoxa ebena is almost entirely black
and £. formosana almost entirely orange
yellow. The hindbasitarsus is extremely
flat, thin, and concave on its outer surface,
similar to that of E. formosana. Because of
the fragility of the specimen, the lancet
was not examained; however, from the
small apical part visible the serrulae ap-
pear to be rather deep similar to E. for-
mosana (Fig. 9).

Eusunoxa formosana Enslin

(Figs. 9, 13, 14, 21)

Eusunoxa formosana Enslin 1911: 99. — Wei 1997:
88 (in subg. Eusunoxa; in key).

Female. — Length, 7.5-8.0 mm. Antenna,
head, thorax, and abdomen yellowish or-
ange, with antennal segments 5-9, apical
third to half of hind femur, and most of
hind tarsus black; apical segments of hind
tarsus paler, more dark orange. Wings yel-
low hyaline, costa and subcosta yellowish,
stigma brownish with margins yellowish,
rest of venation dark brown. Antenna fi-
liform, very slightly incrassinate in mid-
dle; antennal length 1.7X head width; 3rd
segment 1.1 X length of 4th; pedicel as
long as its apical width; scape slightly lon-
ger than its apical width. Clypeus sub-
truncate; malar space distinct but much
less than half diameter of front ocellus;
lower interocular distance 0.8X eye
length. OOL:POL:OCL = 1.00:1.00:0.83,
postocellar area convex, 2X broader than
long at its maximum breadth. Head with
frons densely punctured, punctures deep,

Volume 12, Number 2, 2003

341

close together, separated by narrow ridges
(similar to Fig. 1); postocellar area and
head behind eyes sparsely punctate, shin-
ing. Tarsal claws with inner tooth shorter
than apical one, basal lobe distinct (Fig.
21). Hind basitarsus extremely flat and
concave on its outer surface, 1.7X length
of remaining tarsal segments combined;
4.8 X longer than maximum width. Lancet
in Fig. 9, serrulae deep with almost per-
pendicular anterior margin.

Male. — Length, 7.5 mm. Similar in color
and structure to female. Genitalia in Figs.
13, 14.

Type. — Described from a female from
"Formosa," in the Zoologisches Staats-
sammlung, Miinchen, Germany; not ex-
amined.

Specimens examined. — TAIWAN: Taihor-
in, Formosa, H. Sauter, 7.VI.1911 (1 9,
USNM); Kankau (Koshun) Formosa, H.
Sauter, VI-1912 (1 9, USNM), same except
VII-1912 (1 9, DEI); PingTung Co., Rent-
ing Bot. Garden, subtropical forest, July
10-14, 1980, 260 m, D.R. Davis (1 8,
USNM); Formosa, Sauter, Taihorinaho,
1090.VIII (1 9, BMNH).

Distribution. — Taiwan.

Remarks. — We did not examine the type
since the description and uniqueness of
this species is sufficient for its identity.
This and £. ebena are the only species of
Eusunoxa with a basal lobe on the tarsal
claws and one of the few species (other
than £. ebena, E. punctata and £. buchi) with
a densely punctate frons. The almost en-
tirely orange coloration is also distinctive.
The female ovipositor, especially the
shape of the serrulae (Fig. 9), differs from
other species of Eusunoxa. Male genitalia
(Figs. 13, 14) differ from other species by
the longer parapenis of the genital cap-
sule.

Eusunoxa lissofrons Smith and Saini,
new species

(Figs. 4, 6, 19)

Female. — Unknown.

Male. — Length, 5.5-6.0 mm. Head and

thorax black, abdomen and fore- and mid-
legs brownish, hind leg black. Wings in-
fumated apically from base of stigma, bas-
al half clear; venation dark brown to
black. Antennal length 1.7X head width,
slightly compressed and only feeblv in-
crassinate in middle; pedicel 2X longer
than its apical width; scape as long as
broad at apex; 3rd segment 1.1 X longer
than 4th. Clypeus subtruncate; lower in-
terocular distance 0.9X eye length; malar
space distinct but much less than half di-
ameter of front ocellus; OOL:OCL:POL =
1.00:1.00:0.70; postocellar area convex,
2.8 X broader than long. Head (Fig. 4), tho-
rax, and abdomen smooth, impunctate,
with polished surface. Hind basitarsus
about 1.2X length of remaining tarsal seg-
ments combined, about 4.0 X longer than
maximum width. Tarsal claws without
basal lobe. Genital capsule similar to Fig.
15, penis valve in Fig. 19.

Holotype. — 8, "South India, Anamalia
[Anamalai ?] Hills, Cinchona, 3500 ft., V-
1965, P.S. Nathan" (USNM).

Paratypes. — INDIA: Same data as holo-
type (18, USNM), same except V-1964 (1
8, USNM), X-1967 (2 8, USNM); Coim-
batore, S. India, 9.7.1949, P. Susai Nathan
(1 8 , USNM); Kerala: Permade, Parryar,
3500 ft., 21-9-1995, coll. M.S. Saini (1 8,
PUNJ).

Distribution. — India (Kerala, Tamil
Nadu).

Etymology. — The species name is based
on the shining, impunctate frons.

Remarks. — The impunctate, shining
frons is similar to that of E. ceylonica and
£. nigriceps, but the entirely black colora-
tion precludes the association of E. lissof-
rons because color variation has not been
observed in males of those species. The
male of £. nigriceps has the thorax partly
orange, and the male of £. ceylonica has the
thorax and abdomen mostly orange, un-
like the entirely black thorax and abdo-
men of £. lissofrons. The male penis valve
of £. lissofrons (Fig. 19) is oval and similar
to that of £. ceylonica (Fig. 16); it is less

342

Journal of Hymenoptera Research

similar to the curved valve of £. nigriceps
(Fig. 20). There is no indication that E. lis-
sofrons is a color variant, but, if so, it could
be one of E. ceylonica.

Eusunoxa nigriceps (Rohwer)

(Fig. 20)

Beleses nigriceps Rohwer 1915: 51.
Eusunoxa nigriceps: Smith 1982: 120.— Wei 1997:
89 (in subg. Asunoxa; in key).

Female. — From Rohwer (1915). "Length,
6 mm." "Head and posterior femora be-
yond middle, four posterior tarsi and the
antennae black, the rest of the insect ru-
fous." "Wings distinctly hyaline, venation
dark brown." "Clypeus truncate, surface
coarsely, irregularly punctured;" "front
and posterior orbits shining, impunctate;"
"flagellum gradually thickened until it
reaches the apex of the second joint;" "the
fourth and following joints compressed;
thorax shining, impunctate."

Male (paratype). — Length, 6.2 mm. An-
tenna and head black. Thorax blackish
with tegula, pronotum, V-shaped mark on
mesoprescutum, mesoscutellum, and me-
tanotum dark orange. Abdomen mostly
black. Foreleg orange with tarsus black;
midleg with coxa, trochanter, and femur
orange, tibia and tarsus black; hind leg
with coxa, trochanter, and basal third of
femur orange, apical two-thirds of femur
and entire tibia and tarsus black. Wings
lightly,uniformly infuscated; veins and
stigma black. Antennal length 1.7X head
width; 3rd segment about 1.1 X length of
4th. Lower interocular distance 0.8 X eye
length; postocellar area 2.6 X broader than
long; clypeus subtruncate; malar space lin-
ear; OOL:POL:OCL = 1.00:1.00:0.74. Head
shining, impunctate with very few punc-
tures between antennae and eyes (similar
to Fig. 4). Hind basitarsus 1.5X length of
remaining tarsal segments combined,
about 2.5 X longer than maximum width.
Tarsal claws without basal lobe. Genital
capsule similar to Fig. 15; penis valve in
Fig. 20, markedly curved in lateral view.

Types. — "Described from one female
from Marikuppam, . . . 3,500 feet" and
from "two males, one allotype, from Ban-
galore, . . . 3,000 feet" (South India). Type
and allotype in the Indian Museum, as
stated by Rohwer (1915), but we could not
locate them and they are apparently no
longer in existence. One paratype male is
in the USNM, labeled "Ind. Mus., Banga-
lore, S. India, ca. 3000 ft., 12-X-10, Annan-
dale," "U.S.N.M. Paratype No. 18909."

Distribution. — India (Karnataka).

Remarks. — Rohwer mentioned claws
cleft, with the inner teeth exceeding the
outer. Because the teeth are lateral and the
paratype examined has the inner tooth
shorter, Rohwer probably meant the outer
tooth is longer.

Because we could not examine the ho-
lotype female, we assume Rohwer's asso-
ciation of sexes was correct. The descrip-
tion of the female is based on Rohwer's
(1915) original description. It is very sim-
ilar to the female of £. ceylonica and, so far
as we can determine, can only be separat-
ed by the orange coloration of the tibiae
and hind femur (see preceding key). Sep-
aration of E. nigriceps is based on the para-
type male which, unlike most species and
shared only by E. lissofrons and E. ceylon-
ica, has the head shining and impunctate.
However, the male of E. lissofrons is en-
tirely black, and the male of E. ceylonica
has the thorax and abdomen almost en-
tirely orange. The shape of the male penis
valve (Fig. 20) is markedly different from
those species, and it is based on this fea-
ture that we believe £. nigriceps is a valid
species and not a color variation of the
others. We have not seen specimens that
match Rohwer's description of the female
or the paratype male examined. Thus, we
keep £. nigriceps a separate species until
more material is available for evaluation
of its status.

Eusunoxa punctata Smith and Saini,
new species

(Figs. 1, 8, 11)

Female. — Length, 8.2 mm. Head and an-
tenna black with 1st antennal segment,

Volume 12, Number 2, 2003

343

/>

14

21

Figs. 13-22. 13-20, Male genitalia. 13, Genital capsule, ventral view of left half, ol I usunoxa formosana. 14,
Penis valve, lateral, of E. formosana. 15, Genital capsule, ventral view of left half, of £. ceylonica. 16, Penis valve
of E. ceylonica. 17, Penis valve of E. auricauda. 18, Penis valve of E. semipunctata. 19, Penis valve ol E. lissofrons.
20, Penis valve of E. nigriceps. 21-22, Tarsal claws. 21, E. formosana. 11, E. ceylonica.

area below antenna, clypeus, labrum,
mouthparts, mandible except tip, and very
dim spots lateral to lateral furrows dark
orange. Thorax and abdomen orange, ex-
cept tergites 2-7 dark brownish at center.

Legs orange with midtarsus, apical two-
thirds of hind femur, apical half o\ hind
tibia, and entire hind tarsus black. Wings
hyaline, infumated on apical half; stigma
and costa brownish, rest of venation dark

344

Journal of Hymenoptera Research

brown to black. Antenna subincrassinated
in middle, antennal length 1.7X head
width; pedicel 2X its apical width; scape
as long as its apical width; 3rd and 4th
antennal segments subequal in length.
Clypeus truncate; lower interocular dis-
tance subequal to eye length; malar space
distinct but much less than half diameter
of front ocellus; OOL:POL:OCL = 1.00:
0.96:0.96; postocellar area subconvex, 2.4X
broader than long. Frons densely punc-
tate, punctures large and close together,
separated by narrow ridges (Fig. 1); hind
orbits and postocellar area less punctured,
shining. Hind basitarsus (Fig. 8) 1.7X
length of remaining tarsal segments com-
bined; 3.7X longer than its maximum
width. Tarsal claws without basal lobe.
Lancet as in Fig. 11, with about 21 serru-
lae.

Male. — Unknown.

Holotype. — 9, Nagaland: Chuchuyin-
long, 2800 ft., 9.5.1994, collection M.S. Sai-
ni (PUNJ).

Distribution. — India (Nagaland).

Etymology. — The species name is based
on the densely punctate frons.

Remarks. — In E. punctata, the hind basi-
tarsus is unusually long in comparison to
the length of the remaining tarsal seg-
ments, and the frons is densely punctate,
both character states of which are shared
only with E. formosana and E. ebena. Eu-
sunoxa formosana, however, is almost en-
tirely orange yellow and both £. formosana
and £. ebena have a basal lobe on the tarsal
claws. The shape of the serrulae of the lan-
cets of £. punctata and E. formosana also
differ (Fig. 9, 11). The only other species
with a densely punctate frons and absence
of a basal lobe on the tarsal claws is E.
buchi from the Philippines. However, £.
buchi has a shorter hind basitarsus, only
about 1.4X the length of the remaining
tarsal segments combined, and the basal
two and basal half of the third antennal
segments are dark orange.

Eusunoxa semipunctata
Smith and Saini, new species

(Figs. 2, 7, 12, 18)

Female. — Length, 7.2-8.0 mm. Antenna
and head black. Thorax and abdomen or-
ange, upper half of mesepisternum some-
what more whitish than orange. Legs with
coxae blackish except apices, trochanters
white, femora blackish except extreme ba-
ses orange, fore- and midtibiae and fore-
and midtarsi white with apical 3-4 tarsal
segments infuscated to black. Wings light-
ly infumated apically from base of stigma,
basal halves clear, venation dark brown to
black. Antenna subincrassinated in mid-
dle, antennal length 1.8X head width;
pedicel and scape each 2X longer than
their apical widths; 3rd segment very
slightly longer than 4th. Clypeus subtrun-
cate; lower interocular distance subequal
to eye length; malar space nearly linear;
OOL:POL:OCL = 1.00:0.80:0.68; postocel-
lar area subconvex, 2.9 X broader than
long. Frons with distinct but sparse, wide-
ly separated punctures with broad, shiny
interspaces between them (Fig. 2). Hind
basitarsus (Fig. 7) 1.3X length of remain-
ing tarsal segments combined; 3.9 X longer
than maximum width. Tarsal claws with-
out basal lobe. Lancet as in Fig. 12, with
about 20-21 serrulae.

Male. — Length, 6.5-7.0 mm. Antenna,
head, and thorax black. Color of legs sim-
ilar to that of female. Structure similar to
that of female. Genital capsule similar to
Fig. 15, penis valve in Fig. 18.

Holotype. — 9, "South India, Madras
State. Anamalai Hills, Kadamparai, 3500',
V-'63, P.S. Nathan" (USNM).

Paratypes. — INDIA: Same data as holo-
type (12 9, 43 cT); Nilgiri Hills, Naduva-
tam, S. India, 6000 ft., May 1958, P.S. Na-
than (1 6); South India, Anamalai Hills,
Cinchona, 3500 ft., IV-1969, P.S. Nathan (6
6 ); V-1964 (4 6 ); V-1965 (4 9, 1 6); V-1966
(3 6); V-1967 (4 9,2 c?); IV-1969 (1 6); V-
1968 (2 9, 2 cT); V-1969 (2 9, 2 6); IX-1969,
T.R.S. Nathan (1 6); Poomundi, 1550 m,

Volume 12, Number 2, 2003

345

Kerala, India, 19-9-95 (1 6) (all USNM);
Kerala: Poonmundi Hills, 4800 ft.,
18.5.1995, collection M.S. Saini (1 6,
PUNJ).

Distribution. — India (Kerala).

Etymology. — The name is based on the
presence of widely spaced punctures on
the frons.

Remarks. — Association of sexes is based
on series taken at the same localities and
at the same time. Only the coloration of
the thorax and abdomen differs; the males
black and the females orange. Such color
dimorphism is not unusual in sawflies,
and E. nigriceps apparently also shows col-
or dimorphism. However, males of E. cey-
lonica and E. formosana are similar in color
to the females. Both sexes of E. semipunc-
tata have the distinct but widely spaced
punctures on the frons with shining inter-
spaces (as in Fig. 2). The female is sepa-
rated from those of other species of Eux-
unoxa by the mostly black femora, and the
male by the mostly black legs with the
fore- and midtibiae and tarsi white. The
punctation of the head also separates the
female; no other species with an orange
thorax and abdomen have such dense
punctures. The male is similar to E. auri-
cauda, but the abdomen of E. auricauda is
orange (see remarks under that species).

ACKNOWLEDGMENTS

We thank the following for loan of specimens in
their care: A. Taeger and S. Blank, Deutsches Ento-
mologisches Institut, Eberswalde, Germany; C. Tay-
lor, The Natural History Museum, London, U.K.; I.
Persson, Naturhistoriska Riksmuseet, Stockholm,
Sweden; and E. Kierych, Polska Akademia Nauk,

Warsaw, Poland. Cathy Apgar, Systematic Entomol-
ogy Laboratory, USDA, prepared the photographs.
We thank the following for review of the manuscript:
H. Goulet, Agriculture and Agri-Food Canada, Otta-
wa, and J. W. Brown and E. E. Grissell, Systematic
Entomology Laboratory, USDA, Washington, DC. Fi-
nancial assistance provided by the U.S. Department
of Agriculture (PL-480, Grant No. FG-In-753, Project
No. IN-ARS-418) in collaboration with ICAR, New
Delhi, is also thankfully acknowledged.

LITERATURE CITED

Enderlein, G. 1920 [1919]. Symphytologica II. Zur
Kenntnis der Tenthredininen. Sitzungsberichl der

Gesellschaft Naturforschendcr Freunde zu Berlin, pp.
347-374.

Enslin, E. 1911. Ein Beitrag zur Tenthrediniden-Fauna
Formosas. Societas Entomologica 24: 93-104.

Haris, A. 2000. New Oriental sawflies (Hymenoptera:
Tenthredinidae). Somogyi Muzeumok Kozlemenyei
14: 297-305.

Malaise, R. 1932. A new sawfly from Ceylon (Hym.
Tenthredinidae). Ceylon journal of Science, Section
B 17: 147-148.

Malaise, R. 1963. Hymenoptera Tenthredinoidea, sub-
family Selandriinae, key to the genera of the
world. Entomologisk Tijdskrift 84: 159-315.

Rohwer, S. A. 1915. Some Oriental sawflies in the In-
dian Museum. Records of the Indian Museum 1 1 :
39-53.

Saini, M. S. and J. S. Deep. 1994. First record of Al-
lantinae (Tenthredinidae: Hymenoptera) from
India, journal of the Bombay Natural History Society
91: 47-50.

Smith, D. R. 1982. Symphyta (Hymenoptera) of Sri
Lanka. Proceedings of the Entomological Society of
Washington 84: 117-127.

Togashi, I. 1981. Some sawflies (Hymenoptera, Per-
gidae and Tenthredinidae) from New Britain,
Papua New Guinea, and Palawan Is., Philip-
pines. Kontyti 49: 414-421.

Wei, M. 1997. Review of the genus Eusunoxa with
erection of a new subgenus (Hymenoptera: Blen-
nocampidae). journal of Central South Forestry
University 17: 88-89.

J. HYM. RES.
Vol. 12(2), 2003, pp. 346-354

Larval Anatomy of Orussidae (Hymenoptera)

Lars Vilhelmsen

Zoological Museum, University of Copenhagen, Universitetsparken 15, DK-2100, Denmark,

e-mail: lbvilhelmsen@zmuc.ku.dk

Abstract. — The external and selected parts of the internal anatomy of the larva of Orussus abie-
tinus (Orussidae) are examined The external anatomy is similar to that reported for other Orus-
sidae. The eyes, laterocervicalia, and thoracic legs are absent, and the antennae, maxillae, and
labium are reduced. Each thoracic and abdominal segment has a pair of short transverse rows of
recurved cuticular spines laterodorsally, probably for locomotory purposes. A previously unre-
ported feature is the configuration of the hindgut, which has a set of transverse interlocking
cuticular folds. The mid- and hindgut are anatomically continuous. Overall, the larval anatomy
of Orussidae is highly reduced compared with that of other basal hymenopteran lineages and
resembles that of apocritan larvae. The modifications of the larval anatomy are probably correlated
with the shift in lifestyle from herbivorous to carnivorous, which is less demanding of the sensory,
feeding, and locomotory apparatus.

Orussidae is a small wasp family, com- termediate in Hymenoptera (Hanson and
prising 75 species worldwide (Vilhelmsen Gauld 1995; Vilhelmsen 1997).
in press). Traditionally, the Hymenoptera Nutall (1980) provided a very brief de-
have been subdivided in two suborders, scription of the larva of Guiglia schauins-
the almost exclusively herbivorous 'Sym- landi (Ashmead, 1903), a New Zealand
phyta' and the predominantly entomoph- species. The only detailed descriptions of
agous Apocrita; Orussidae were usually the immature stages of an orussid species
placed in the 'Symphyta' because of the are by Rohwer and Cushman (1917) and
absence of the wasp-waist in the adults, Parker (1935), who dealt with Orussus oc-
the defining feature of the Apocrita. How- cidentalis Cresson, 1879, a Nearctic species,
ever, recent phylogenetic treatments of the The section on Orussidae in Yuasa's (1922)
Hymenoptera unequivocally have re- treatment of non-apocritan larvae is based
trieved Orussidae + Apocrita as an ex- entirely on Cushman and Rohwer (1917).
tremely well supported clade (Ronquist et They noted that the larva of Orussus share
al. 1999, Vilhelmsen 2001). Information on many traits with those of apocritan Hy-
the biology of most species of Orussidae menoptera (eyes, most head appendages,
is scarce or non-existent; what evidence and thoracic legs very reduced), whereas
there is (see Vilhelmsen et al. 2001) indi- the anatomy of adult Orussidae apparent-
cates that the larvae are ectoparasitoids of ly resembles that of 'Symphyta'. This mix
woodboring insect larvae, usually Bupres- of features and the parasitoid lifestyle led
tidae (Coleoptera). This was probably the Rohwer and Cushman (1917) to erect a
lifestyle of the common ancestor of all par- new suborder, the Idiogastra, comprising
asitoid Hymenoptera. The position of the just the Orussidae.

woodboring 'siricoid' superfamilies as Parker (1935) observed a few internal

successive outgroups to the Orussidae- features of the head anatomy of O. occi-

Apocrita clade indicates that the parasit- dentalis, but otherwise information about

oid lifestyle evolved via a woodboring in- the internal larval anatomy of the family

Volume 12, Number 2, 2003 347

is wanting. Orussidae were not included lected in a kerosene-acetic acid-dioxane

in the survey of 'symphytan' larvae by solution (KAAD). Only a limited number

Maxwell (1955). Specifically, it is not of observations of the external anatomy

known whether there is any connection could be carried out on these specimens.

between the mid- and hindgut. The sepa- External features.— Initial examination of

ration of these gut sections and the post- external features on specimens kept in eth-

ponement of voiding of the gut contents anol was carried out with a Leica MZ Apo

(as a meconium) until just prior to pupa- dissection microscope.

tion is a near universal feature of the Apo- SEM.— Specimens preserved in ethanol

crita (Hanson and Gauld 1995; Quicke were critically point dried and coated with

1997); it usually is interpreted as an im- Pt/Pd in a Jeol JFC-2300HR high resolu-

portant adaptation to the parasitoid life- tion fine coater. They were examined with

style, especially endoparasitism, apparent- a Jeol JSM-6335F field emission scanning

ly serving to prevent contamination of the electron microscope.

host. However, the condition in Orussidae Histological sections. — Specimens pre-

and some of the putatively basal apocritan served in ethanol or Pampels fluid were

taxa (Stephanidae, Megalyridae) has not dehydrated in butanol and embedded in

been investigated, making it impossible to paraplast. Sagittal sections of 8-12 fxm

deduce exactly when the feature arose in thickness, depending on the size of the

the evolutionary history of parasitic Hy- specimen, were cut with a microtome. The

menoptera. sections were treated in a trichrome stain

In the present study, I investigate the (Weigert's haematoxylin-bluish erythro-

larval anatomy of two species of Orussi- sin-fast green, preceded by phosphomo-

dae in some detail. In addition to the ex- lybdic acid); permanent preparations were

ternal features already described by Roh- made in entellan.

wer and Cushman (1917), selected internal

features are examined. The findings are KhSULIS

discussed in relation to features observed The descriptions are based entirely on

in other hymenopteran larvae. the prepupae and early instar of O. abie-

timts. The observations that could be

MATERIALS AND METHODS made on the G sd„mm.ll„ll1i specimens

Larvae of Orussus abietimis (Scopoli, did not reveal any significant differences

1763) and Guiglia schauinslandi (Ashmead, between the two species.

1903) were available for study. Unfortu- Overall. — Body elongate, dorsoventrally

nately, only the material of the former was flattened (less conspicuous in early instar).

in good condition, having been collected Body outline in cross-section with distinct

recently (1998 or later) and preserved in bend laterally between upper and lower

80% ethanol or Pampels fluid and subse- surface. Head hypognathous, boundary

quently transferred to ethanol. In total, between head and prothorax indistinct in

five larvae of O. abietinus were examined, early instar, especially laterally. Segment

Of these, four were final instars or pre- boundaries otherwise distinctly demarcat-

pupae, collected in early spring prior to ed by deep furrows dorsally, especially on

pupation, and one was an early instar col- abdomen. Body cuticle wrinkled, but

lected later in spring on a moldy pupa of without regular transverse annuli or other

Buprestis sp. (Coleoptera, Buprestidae). regular intrasegmental subdivisions.

Four larvae of G. scliaiiiiislandi were ex- Small trichoid sensilla scattered over body

amined. All were in rather poor condition, surface. All thoracic and abdominal seg-

having been preserved in 95% ethanol for ments dorsally with paired, transverse

40+ years after having initially been col- rows of 2—1 (early instar) or 3-5 (prepupa)

348

Journal of Hymenoptera Research

Figs. 1-6. Orussus abietinus, head anatomy. 1, 3-6. Prepupa. 1, Head, ventral view, arrow indicates position
of anterior tentorial pit: an = antenna; la = labium; lb = labrum; mx = maxilla. 2, Early instar larva, mouth-
parts, lateral view: md = mandible (only base visible). 3, Antenna. 4, Tips of mandibles, ventral view. 5,
Maxilla, arrows indicate trichoid sensilla (two leftmost broken). 6, Labium, arrows indicate trichoid sensilla:
so = salivarial orifice.

sligthly recurved cuticular spines subme-
dially (Figs. 7-10).

Head. — Head capsule weakly sclero-
tized. Eye absent. Antenna one-segment-
ed, short, rounded, situated lateroventral-
ly on head capsule (Figs 1-2), distally with
two short peglike sensilla in shallow de-
pression (Fig. 3). Clypeus weakly delim-
ited dorsally; anterior tentorial pits faintly
visible as shallow, elongate depressions

(Fig. 1). Circular cuticular structure pre-
sent medially of anterior tentorial pits, not
visible externally. Small, transversely
elongate sclerite present between clypeus
and labrum. Labrum fleshy, slightly bifid
apically, with 7-8 trichoid sensilla antero-
lateral^. Mandibular base broad, fleshy
(Fig. 2), mandibular articulations dis-
placed medially, not visible externally on
head capsule. Distal part of mandible slen-

Volume 12, Number 2, 2003 349

der and heavily sclerotized, arising medi- surrounded by muscular sheath, except
ally from base, partly covered by labrum for part posteriorly of cuticular folds (Fig.
anteriorly and maxilla posteriorly (Fig. 2), 12); sheath with transverse and outer Ion-
terminating in three distinct cuticular gitudinal fibers. Prepupa similar to early
teeth visible ventrally (Fig. 4). Maxilla un- instar except for following: Midgut with
differentiated, transversely elongate fleshy straight walls, apparently empty except
lobe, anteriorly abutting labrum; maxil- for peritrophic membrane. Boundary be-
lary palp not developed, three trichoid se- tween mid- and hindgut marked by one
tae situated in middle of maxillar lobe pair of malphigian tubules opening into
(Fig. 5). Labium narrow anteriorly, broad- lateral part of gut. Epidermis retracted
ening posteriorly (Fig. 6), weakly delimit- from cuticle in posteriormost abdominal
ed from maxilla by shallow oblique sul- segment (Fig. 13), interspace between epi-
cus, retracted relative to maxillae. Salivar- dermis and cuticle occupied by amor-
ial orifice transverse slit surrounded by phous material traversed by elongate tis-
sclerotized cuticle (barely discernible in sue strands. Epidermis of anterior part of
early instar larva), situated subapically on hindgut not thickened. Muscle sheath not
labium. Labial palp not developed, paired extending very far along hindgut, not
short trichoid sensilla present posterolat- reaching region of cuticular folds, com-
erally. paratively less developed than in early in-

Thorax. — Laterocervicalia absent. Ante- star larva. Hindgut walls with 18-20 close-

rior thoracic spiracle well developed, ap- ly appressed cuticular folds (Fig. 14); folds

parently situated slightly anterior to transversely oriented medially, obliquely

boundary between pro- and mesothorax. posteromedially oriented laterally.
Posterior thoracic spiracle vestigial, rep-

resented by small, elongate sclerotisation ui^u^iui\

just posterior to meso/metathoracic The larva of O. abietinus closely resem-

boundary. Thoracic legs entirely absent. bles O. occidental is (see Rohwer and Cush-

Abdomen. — Ten segments present. Seg- man 1917) in all external features; this is

ments 1-8 with well developed spiracles hardly surprizing, since the two species

situated laterodorsally (Fig. 7); spiracles are probably sister taxa (Vilhelmsen in

circular, rim well sclerotized. Abdominal press). The more distantly related G.

legs, suranal process, suranal hook, and schauinslandi does not differ significantly

subanal appendages absent. Anal slit sit- from Orussus spp. in the features that

uated posteriorly on segment 10, trans- could be observed. However, all three spe-

versely elongate (Figs 9, 10), shallow (can- cies are comparatively derived within

not be pried open with a fine needle). Orussidae, making inferences about the

Hindgut. — Posterior part of midgut of ground plan states for the family tenuous.

early instar larva with convoluted walls. The larval head anatomy of Orussidae

Gut contents amorphous, with dark is highly reduced compared with other

grains. Malphigian tubules not observed, non-apocritan Hymenoptera. The eye is

Mid- and hindgut continuous (Fig. 11). entirely absent (a condition also observed

Hindgut lined with unsclerotized cuticle in 'siricoid' superfamilies), the antenna is

becoming progressively thicker posterior- one-segmented and of the mouthparts,

ly. Anterior part of hindgut with some- only the mandibles appear to be function-

what thickened epidermis. Hindgut with al, both the maxilla and labium having

walls straight, except for 10-12 narrow lost all traces of endites and palps and be-

transverse cuticular folds posteriorly (Figs ing immovable relative to the head cap-

11, 12); cuticular folds correspond to con- sule. In contrast, most 'Symphyta' have

cavities in opposite wall. Most of hindgut multi-segmented antenna as well as dif-

350

Journal of Hymenoptera Research

ferentiated endites and palps on the max-
illa and labium (see Yuasa 1922). The
mandibles of herbivorous 'Symphyta' are
usually well sclerotized throughout with
external articulations and broad shearing
teeth distally. The condition of the man-
dibles (partly concealed by labrum and
maxilla, proximal part unsclerotized, dis-
tal part narrowed, see also Rohwer and
Cushman 1917, fig. 2a) observed in Orus-
sidae is apparently similar to that of
Schlettererius cinctipes (Cresson) (Stephan-
idae; Short 1978, fig. 20; Taylor 1967). Gen-
erally, the head anatomy of orussid larva
is much closer to that of Apocrita than of
other 'Symphyta'. With regard to maxillar
and labial palps, those of Orussidae are
even more reduced than some Apocrita
(e.g., some Ichneumonoidea (Finlayson
1987) and Aculeata (Evans 1987; McGinley
1987)), which have papilliform, one-seg-
mented palps.

The thoracic and abdominal anatomy of
the larvae of Orussidae and Apocrita are
also reduced relative to more basal Hy-
menoptera. The laterocervicalia, which in
representatives of most 'symphytan' su-
perfamilies articulates with the head cap-
sule anteriorly and connects to the pro-
notum posteriorly (Vilhelmsen unpubl.),
is entirely absent in Orussidae (see also
Parker 1935) and apparently also in Apo-
crita, providing a closer integration of the
head and thorax in the two latter taxa. The
thoracic legs are entirely absent, having
become progressively reduced within the
basal lineages of the Hymenoptera (Vil-
helmsen 2001). A suranal process, a heavi-
ly sclerotized projection on the 10th ab-
dominal segment dorsally of the anal slit
prominent in Cephidae, Anaxyelidae, Sir-
icidae, and Xiphydriidae (see Yuasa 1922,
pi. xii) is absent from Orussidae and Apo-
crita. The transverse rows of cuticular
spines dorsally have been reported only
from Schlettererius cinctipes (Stephanidae;
Taylor 1967) outside the Orussidae, al-
though in this species, they are only pre-
sent in the prepupa. Their function in

orussid larvae is perhaps to facilitate
movement along wood galleries, in-
creased leverage being provided by the
deep dorsal furrows between the body
segments. Cooper (1953) observed ovipo-
sition by female Orussus in wood galleries
some distance from potential hosts; he in-
terpreted this as indicating that the larvae
might be frass feeders. Alternatively, the
early instar larva might actively seek out
the host within its galleries, if the female
was unable to reach the host with its ovi-
positor; this possibility was considered
but rejected by Cooper.

Given the well corroborated monophyly
of Orussidae + Apocrita, the interpreta-
tion of the phylogenetic significance of the
shared reductional features in the larval
anatomy is fairly straightforward: they
represent synapomorphies for these two
taxa. Some of these characters have al-
ready been included in recent phylogenet-
ic treatments of the Hymenoptera (e.g.,
number of larval antennal segments, re-
duction of larval thoracic legs, characters
224 and 228 in Vilhelmsen 2001), whereas
as others (reduced maxillar and labial ap-
pendages, absence of laterocervicalia)
have been ignored until now.

The functional significance of the ana-
tomical modifications can be interpreted
in the context of the shift in larval lifestyle
from herbivorous/woodboring to carniv-
orous/parasitoid having taken place in
the common ancestor of Orussidae and
Apocrita. The reduced sensory capabilities
(loss of eyes, reduction of antennae) reflect
the confined habitat (galleries in wood) of
the larvae of the earliest parasitoid wasps
and their closest relatives, the woodwasps.
The reduction of the mouthparts is per-
haps a consequence of the shift in food
source from particulate plant matter that
requires considerable handling and chew-
ing to a much less demanding diet of in-
sect body fluids. Once an ectoparasitoid
wasp larva is in contact with its host, it is
required to do little more than puncture
the integument of the latter to gain access

Volume 12, Number 2, 2003

351

Figs. 7-10. Orussus abietinus, abdominal anatomy. 7-8, Early instar larva, posterior to the left. 7, Dorsal part
of abdominal segment: arrows indicate cuticular spines; sp = spiracle. 8, Recurved cuticular spines. 9-10,
Prepupa. 9, 10th abdominal segment, posterior view, arrows indicate cuticular spines. 10, Same, lateral view,
arrow indicates anal slit.

352

Journal of Hymenoptera Research

Figs. 11-14. I Orussus abietinus, internal anatomy of hindgut (posterior to left). 11-12. Early instar larva. 11,
Overview of hindgut region, arrow indicates transition between mid- and hindgut: mus = muscle sheath. 12,
Details of cuticular folds in hindgut. 13-14, Prepupa. 13, Overview of hindgut region, arrow indicates tran-
sition between mid- and hindgut, rectangle indicates approximate extent of Fig. 14: epi = retracted epidermis.
14, Details of cuticular folds in hindgut.

to its contents, a purpose for which the
narrow, pointed mandibles of the orussid
larva probably is admirably suited. The
reduction of the locomotory apparatus
again reflects the confined larval habitat,
being taken even further than in the Ce-
phidae and woodwasp families, which re-
tain vestiges of thoracic legs. The suranal
process present in these taxa is also absent
in Orussidae and Apocrita, a change that
perhaps is correlated with the shift in food
source. This feature is needed as a brace
by the endophytophagous taxa when
chewing a tunnel through tough plant ma-
terial; the larvae of Orussidae and wood-
living Apocrita do not chew their own
tunnels and hence do not need a posterior
brace.

The configuration of the hindgut in the
early instar larva and prepupa of Orussus
is unlike anything else reported from Hy-
menoptera. The cuticular folds matching
concavities in the opposite walls and the
extension of the muscular sheath sur-
rounding the hindgut to include the part
with the folds indicate that even though
the mid- and hindgut are not anatomically
separated, the larva is able to close the
hindgut by muscular action. In the pre-
pupa, the opposite walls of the hindgut
abut in the region with the cuticular folds
even though the muscular sheath does not
extend this far back; this may be caused
by the loosening of the prepupal cuticle
and the retraction of the epidermis and as-
sociated musculature prior to pupation.

Volume 12, Number 2, 2003

353

The connection between mid- and hind-
gut in 'Symphyta' whose embryology
have been examined (e.g., Pontania caprae
Linnaeus (Tenthredinidae), see Ivanova-
Kasas 1959) is established already at the
time of hatching from the egg. The ana-
tomical separation caused by the failure of
the two gut sections to join during the em-
bryological development (Hanson and
Gauld 1995) has been reported from a
wide range of apocritan superfamilies:
Ceraphronoidea (Megaspilidae: Dendroce-
rus (= Lygocerus) spp., see Haviland 1920a,
fig. 14), Chalcidoidea (Pteromalidae: Spa-
langia muscidarum Richardson, see Rich-
ardson 1922, fig. 7; Asaphes vulgaris Walk-
er, Pachycrepis clavata Walker, see Havi-
land 1922), Cynipoidea (Figitidae: Charips
spp., see Haviland 1920b, fig. 11a), Ichneu-
monoidea (Ichneumonidae: Pimpla turi-
onellae (Linnaeus), see Fiihrer and Willers
1986, fig. 3), Platygastroidea (Platygastri-
dae spp., see Marchal 1906, pi. xviii: 24),
Proctotrupoidea (Proctotrupidae: Phaenos-
erphus viator (Haliday), see Eastham 1929,
fig. 12). It is often accompanied by consid-
erable differentiation between hindgut re-
gions (epidermis and muscle layer of var-
iable thickness, formation of valves; e.g.,
Eastham 1929, Fiihrer and Willers 1986,
Haviland 1920b) that is less conspicuous
in O. abietinus. However, many 'Symphy-
ta' also have the hindgut differentiated
into several regions (Maxwell 1955).

The functional and phylogenetic signif-
icance of the larval hindgut anatomy of
Orussus is difficult to interpret without
further information about the orussid life-
style and investigation of larvae of some
of the basalmost apocritan ectoparasitoid
taxa (e.g., Stephanidae and Megalyridae).
It is possible, but entirely conjectural, that
the cuticular folds help the orussid larva
to clamp its hindgut shut for most of its
development, thus preventing contami-
nation of its host. However, the expulsion
of a meconium prior to pupation as seen
in most apocritans examined was not ob-
served in a couple of hatchings of O. abie-

tinus (Vilhelmsen unpubl.). The anatomi-
cal position (posteriorly in the hindgut) of
the cuticular folds in Orussus is not ho-
mologous with the position (boundary be-
tween mid- and hindgut) of the separation
of the gut sections in Apocrita, making it
unlikely that the latter evolved from the
former. Thus, this study has revealed an-
other intriguing feature occurring in par-
asitic Hymenoptera, rather than elucidat-
ing the evolution of an already known
trait.

ACKNOWLEDGMENTS

Hans Ahnlund, Gnesta, Sweden, provided invalu-
able assistance with collecting material of O. abietinus.
Toni Withers, Forest Research Institute, Rotorua,
New Zealand, graciously made the material of G.
schauinslandi available for study. Dave Smith and Eric
Grissell, both in Systematic Entomology Laboratory,
Washington, USA commented on an earlier version
of the manuscript.

LITERATURE CITED

Cooper, K. W. 1953. Egg gigantism, oviposition, and
genital anatomy: their bearing on the biology
and phylogenetic position of Orussus (Hymenop-
tera: Siricoidea). Proceedings of the Rochester Acad-
emy of Sciences 10: 38-68.

Eastham, L. E. S. 1929. The post-embryonic develop-
ment of Phaenoserphus viator Hal. (Prototrypoi-
dea), a parasite of the larva of Pterostichus niger
(Carabidae), with notes on the anatomy of the
larva. Parasitology 21: 1-21 + 3 plates.

Evans, H. E. 1987. Families: Bethylidae, Dryinidae,
Chrysididae, Scoliidae, Tiphiidae, Mutillidae,
Sapygidae, Formicidae, Rhopalosomatidae, Pom-
pilidae, Vespidae, Sphecidae. Pp. 670-688 //;:
Stehr, F.W. Immature Insects. Kendall/Hunt Pub-
lishing Company, Dubuque, Iowa, USA, 754 pp.

Finlayson, T. 1987. Ichneumonoidea. I'p. 649-663 /;;:
Stehr, F.W. Immature Insects. Kendall/Hunt Pub-
lishing Company, Dubuque, Iowa, USA, 7^4 pp.

Fiihrer, E. and D. Willers. 1986. The anal secretion oi
the endoparasitic larva Pimpla lurionellae: Sites of
production and effects, lourual of Inset I Physiology
32: 361-367.

Hanson, P. E and I. D. Gauld. L995. The Hymenoptera
of Costa Rica. Oxford University Press, Oxford,
893 pp.

Haviland, M. D. 1920a. On the bionomics and devel-
opment of Lygocerus testaceimanus and Lygocerus
cameroni Kieffer (Proctotrypoidae-Ceraphroni-
dae), parasites of Aphidius, Braconidae. Quarterly
Journal of microscopical Science 65: 101 127.

354

Journal of Hymenoptera Research

Haviland, M. D. 1920b. On the bionomics and post-
embryonic development of certain cynipid hy-
perparasites of aphides. Quarterly Journal of mi-
croscopical Science 65: 451^478.

Haviland, M. D. 1922. On the post-embryonic devel-
opment of certain chaicids, hyperparasites of
aphides. Quarterly Journal of microscopical Science
66: 321-338.

Ivanova-Kasas, O. M. 1959. Die embryonale Entwick-
lung der Blattwespe Pontania caprae L. (Hyme-
noptera, Tenthredinidae). Zoologische Jahrbucher,
Abteilung fiir Anatomie und Ontogenie der Tiere 77:
193-228.

Marchal, P. 1906. Recherces sur la biologie et le de-
velopement des hymenopteres parasites. Les pla-
tygasters. Archives de zoologie experimental et ge-
nerate Serie 4, 4: 475-640 + 8 plates.

Maxwell, D. E. 1955. The comparative internal larval
anatomy of sawflies (Hymenoptera: Symphyta).
The Canadian Entomologist 87, supplement 1, 132

PP-

McGinley, R. J. 1987. Families: Colletidae, Oxaeidae,

Halictidae, Andrenidae, Mellitidae, Megachili-
dae, Anthophoridae, Apidae. Pp. 689-704 In:
Stehr, F.W. Immature Insects. Kendall/Hunt Pub-
lishing Company, Dubuque, Iowa, USA, 754 pp.

Nutall, M. J. 1980. Insect Parasites of Sirex. Forest and
Timber Insects in New Zealand 47: 12 pp. (unpa-
ginated).

Parker, H. L. 1935. Note on the anatomy of tenthre-
dinid larvae with special reference to the head.
Bollcttino del Laboratorio di Zoologia Generale e
Agraria delta R. Scuola Superiore d'Agricoltura in
Portici 28: 159-191.

Quicke, D. L. J. 1997. Parasitic Wasps. Chapman and
Hall, London, 470 pp.

Richardson, E. 1922. Studies on habits and the devel-
opment of a hymenopterous parasite, Spalangia

muscidarum Richardson. Journal of Morphology 24:
513-557.

Rohwer, S. A. and R. A. Cushman. 1917. Idiogastra,
a new suborder of Hymenoptera with notes on
the immature stages of Oryssiis. Proceedings of the
Entomological Society of Washington 19: 89-98.

Ronquist, F., A. P. Rasnitsyn, A. Roy, K. Erikson, and
M. Lindgren. 1999. Phylogeny of the Hymenop-
tera: A cladistic reanalysis of Rasnitsyn's (1988)
data. Zoologica Scripta 28: 13-50.

Short, J. R. T. 1978. The final larval instars of the Ich-
neumonidae. Memoirs of the American Entomolog-
ical Institute 25, 508 pp.

Taylor, K. L. 1967. Parasitism of Sirex noctilio F. by
Schlettererius cinctipes (Cresson) (Hymenoptera:
Stephanidae). Journal of the Australian Entomolog-
ical Society 6: 13-19.

Vilhelmsen, L. 1997. The phylogeny of lower Hyme-
noptera (Insecta), with a summary of the early
evolutionary history of the order. Journal of Zoo-
logical Si/stematics and evolutionary Research 35:
49-70.

Vilhelmsen, L. 2001. Phylogeny and classification of
the extant basal lineages of the Hymenoptera (In-
secta). Zoological Journal of the Linnean Society 131:
393-442.

Vilhelmsen, L. in press. Phylogeny and classification
of the Orussidae (Insecta: Hymenoptera), a basal
parasitic wasp taxon. Zoological Journal of the Lin-
nean Society.

Vilhelmsen, L., N. Isidoro, R. Romani, H. H. Basibu-
yuk, and D. L. J. Quicke. 2001. Host location and
oviposition in a basal group of parasitic wasps:
the subgenual organ, ovipositor apparatus, and
associated structures in the Orussidae (Hyme-
noptera, Insecta). Zoomorphology 121: 63-84.

Yuasa, H. 1922. A classification of the larvae of Ten-
thredinoidea. Illi>wis Biological Mo)iographs 7(4),
172 pp.

J. HYM. RES.
Vol. 12(2), 2003, pp. 355-357

NOTE

First Report of Male Sleeping Aggregations

in the Pollen Wasp Celonites abbreviatus (Villers, 1789)

(Hymenoptera: Vespidae: Masarinae)

F. Amiet and V. Mauss

(FA) Forststr. 7, CH-4500 Solothurn, Switzerland;

(VM) Staatliches Museum fur Naturkunde, Abt. Entomologie, Rosenstein 1,

D-70191 Stuttgart, Germany, email: volker.mauss@stechimmenschutz.de

The nightly resting behaviour of solitary
and social bees, and probably also of
many other Aculeata, constitutes a sleep-
like state with many neurophysiological
parallels to mammalian sleep (Kaiser and
Steiner-Kaiser 1983, Kaiser 1988, 1995).
Sleeping behaviour has been well docu-
mented in many species of solitary wasps
(O'Neill 2001: 294). Interspecific variation
of sleep is evident in (1) the location of the
sleeping site, (2) the postures adopted
during sleep, and (3) whether the wasp
sleeps in the company of other members
of its own species, members of the oppo-
site sex, and members of other species
(O'Neill 2001: 294). In contrast, present
knowledge of sleeping behaviour of male
pollen wasps is poor. Males of some spe-
cies of the genus Ceramius seek overnight
shelter in conspecific nests that may or
may not contain females (Brauns 1910,
Gess 1996: 63, Mauss 1996). Celonites andrei
Brauns was observed to spend the night
sleeping on vegetation (Brauns 1910), and
a male of Celonites abbreviatus slept curled
up around a blade of grass (Bischoff 1927:
Fig. 29). Males of Masarina mixta Richards
were commonly observed to sleep in bell-
shaped flowers of Wahlenbergia (Campan-
ulaceae) on which they and the females
forage during the daytime (Gess 1996: 63).

Male sleeping aggregations have not
previously been recorded for any species
of the Masarinae (Gess 1996: 63), although

they have been observed for numerous
other wasps and bees (summarised by Bis-
choff 1927: 62-64, Westrich 1989: 124,
O'Neill 2001, Wcislo 2003). We describe
here for the first time, male sleeping ag-
gregations in C. abbreviatus which were
discovered at two different locations in
Switzerland.

(1) Pfynwald (07°35'E 46°38'N), Rhone
Valley, Wallis, Switzerland, July 1977, obs.
F. Amiet, A. Krebs: In the evening four
males were sleeping in an aggregation on
a withered, branched stem of an herba-
ceous perennial plant about 0.2 m above
the ground. A photograph of this aggre-
gation was reproduced in Witt (1998: 192
bottom right), however it was not further
commented on, and it was erroneously
shown in the vertical position. Distance
between the males varied from 0 (i.e., in
physical contact with each other) to about
8 mm. All males adopted the same typical
posture: They curled their bodies around
the stem so that the tip of their metasoma
covered the ventral part of the clypeus.
Antennae and legs were pulled up under
the mesosoma, and the wings were folded
underneath the metasoma (Figs. 2, 3, the
identical posture as in Bischoff 1927, fig.
29). The males were observed to aggregate
and sleep on the same stem on several
consecutive nights.

(2) Berner Oberaland near Boltigen
(07°22'E 46°38'N), Simmental, Switzer-

356

Journal of Hymenoptera Research

Figs. 1-3. Celonites abbreviates, male sleeping aggregation. 1, Berner Oberaland, August 1997 (actual body
length of males about 8 mm). 2, Sleeping posture (viewed from left), Pfynwald, July 1977 (Photo: A. Krebs).
3, Sleeping posture (viewed from ventral), Pfynwald, July 1977.

Volume 12, Number 2, 2003

357

land, at the foot of a scree slope, August
1997, obs. F. Amiet: The sky was cloudy
and whenever the sun became obscured
by a larger cloud some males of C. abbrev-
iatus alighted on two withered stems of
herbaceous perennial plants about 0.25 m
above the ground (Fig. 1) and adopted the
typical sleeping posture. Moments later
when the sun emerged again some of
these males became active and flew away.
From three o'clock p.m. (Central Europe-
an Time) onward all the males remained
in the sleeping posture on the stems al-
though the sun still shined intermittently.
At four o'clock p.m. it commenced to rain.
A maximum of 14 males were observed
on both stems. Distance between the
sleeping males in the aggregation ranged
from 0 to about 8 mm.

Other clades of the Vespidae in which
male sleeping aggregations occur are the
Euparagiinae (Euparagia scutellaris Cres-
son, Moore 1975) and at least six genera
of the Eumeninae (Labus, Bischoff 1927: 62,
and Ancistrocerus, Eumenes, Pterocheihis,
Rhynchium, Stenodynerus, Linsley 1962).
The sleeping postures of the eumenine
wasps studied differ distinctly from that
in Celonites abbreviatus: Males of the Eu-
meninae attach themselves to the sub-
strate with their legs and mandibles (Lin-
sley 1962). Their wings are folded but ex-
tend outward at an angle of approximate-
ly 45 degrees. Unfortunately, the sleeping
posture of Euparagia cannot be brought
into context since it was not described in
sufficient detail by Moore (1975). The evo-
lutionary significance of aggregated sleep-
ing is uncertain (Evans et al. 1986), but it
has been suggested that it offers protec-
tion against predators, may influence ther-
moregulation or may be associated with

mating behaviour (cf. Freeman and John-
ston 1978).

LITERATURE CITED

Bischoff, H. 1927. Biologic der Hymenopteren. Springer
Verlag, Berlin. 598 pp.

Brauns, H. 1910. Biologisches iiber siidafrikanische
Hymenopteren. Zeitschrift fiir wissenschaftliche In-
sekten Biologic 6: 384-387, 445-447.

Evans, H.E., O'Neill, K.M. and O'Neill, R.P. 1986.
Nesting site changes and nocturnal clustering in
the sand wasp Bembecinus quinquespinosus (Hy-
menoptera: Sphecidae). journal of the Kansas En-
tomological Society 59: 280-286.

Freeman, B.E. and Johnston, B. 1978. Gregarious
roosting in the sphecid wasp Sceliphron assimile.
Annals of the Entomological Society of America 71:
435-441.'

Gess, S.K. 1996. The pollen wasps — Ecology and natural
history of the Masarinae. Harvard University Press,
Cambridge, Massachusetts. 340 pp.

Kaiser, W. 1988. Busy bees need rest, too. Behavioural
and electromyographical sleep signs in honey-
bees. Journal of comparative physiology A 163: 565-
584.

Kaiser, W. 1995. Rest at night in some solitary bees —
a comparison with the sleep-like state of honey
bees. Apidologie 26: 213-230.

Kaiser, W. & Steiner-Kaiser, J. 1983. Neuronal corre-
lates of sleep, wakefulness and arousal in a di-
urnal insect. Nature 301: 707-709.

Linsley, E.G. 1962. Sleeping aggregations of aculeate
Hymenoptera. Annals of the Entomological Society
of America 55: 148-164.

Mauss, V. 1996. Contribution to the bionomics of Cer-
amius tuberculifer Saussure (Hymenoptera, Ves-
pidae, Masarinae). journal of Hymenoptera Re-
search 5: 22-37.

Moore, W.S. 1975. Observations on the egg laying
and sleeping habits of Euparagia scutellaris Cres-
son (Hymenoptera, Masaridae). The Pan-Pacific
Entomologist 51: 286.

O'Neill, K.M. 2001. Solitary wasps. Cornell LTniversir)
Press, Ithaca, London. 406 pp.

Wcislo, W.T. 2003. A male sleeping roost of a sweat
bee, Augochlorella neglectula (Ckll.) (Hymenop-
tera: Halietidae), in Panama, journal of the Kansas
Entomological Society 7b: 55-59.

Westrich, P. 1989. Die Wildbienen Baden-Wurttembergs.
Eugen Ulmer Verlag, Stuttgart. 972 pp.

Witt, R. 1998. Wespen beobachten, bestimmen. Natur-
buch Verlag, Augsburg. 360 pp.

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Microsoft Word). Authors should keep backup copies of all material sent to the Editor. The Society cannot be
responsible for diskettes or text mislaid or destroyed in transit or during editing.

Illustrations should be planned for reduction to the dimension of the printed page (14 X 20.5 cm, column
width 6.7 cm) and allow room for legends at the top and bottom. Do not make plates larger than 14 X 18 in.
(35.5 X 46 cm). Individual figures should be mounted on a suitable drawing board or similar heavy stock.
Photographs should be trimmed, grouped together and abutted when mounted. Figure numbers should be
on the plate. Include title, author(s) and address(es), and illustration numbers on back of each plate. Original
figures need not be sent until requested by the editor, usually after the manuscript has been accepted. Reference to
figures /tables in the text should be in the style "(Fig. 1)" "(Table 1)". Measurements should be in the metric
system.

Electronic plates may be submitted on disc or via email. They must be fully composited, labeled, and
sized to fit the proportions of the journal page. Line art should be scanned at 1200 dpi (minimum input reso-
lution is 600 dpi). Color or grayscale (halftone) images should have a dpi of 300-350. Color files should be in
CMYK and not RGB. Graphics should be submitted as TIFF or EPS files. No PowerPoint or Word/Word Perfect
files with images embedded in them are acceptable.

All papers must conform to the International Code of Zoological Nomenclature. The first mention of a plant
or animal should include the full scientific name including the authority. Genus names should not be abbre-
viated at the beginning of a sentence. In taxonomic papers type specimens must be clearly designated, type
depositories must be clearly indicated, and new taxa must be clearly differentiated from existing taxa by
means of keys or differential diagnoses. Authors are required to deposit all type material in internationally
recognized institutions (not private collections). Voucher specimens should be designated for specimens
used in behavioral or autecological studies, and they should be deposited similarly.

Acceptance of taxonomic papers will not require use of cladistic methods; however, authors using them
will be expected to specify the phylogenetic program used (if any), including discussion of program options
used. A data matrix should be provided if the subject is complex. Cladograms must be hung with characters
and these should include descriptors (not numbers alone) when feasible. The number of parsimonious
cladograms generated should be stated and reasons given for the one adopted. Lengths and consistency
indices should be provided. Adequate discussions should be given for characters, plesiomorphic conditions,
and distributions of characters among outgroups when problematical.

References in the text should be (Smith 1999), without a comma, or Smith (1999). Two articles by a single
author should be (Smith 1999a, 1999b) or Smith (1999a, 1999b). For multiple authors, use the word "and,"
not the symbol "&" (Smith and Jones 1999). For papers in press, use "in press," not the expected publication
date. The Literature Cited section should include all papers referred to in the paper. Journal names should
be spelled out completely and in italics.

Charges. Publication charges are $10.00 per printed page. At least one author of the paper must be a
member of the International Society of Hymenopterists. Reprints are charged to the author and must be
ordered when returning the proofs; there are no free reprints. Author's corrections and changes in proof are
also charged to the author. Color plates will be billed at full cost to the author.

All manuscripts and correspondence should be sent to:

Dr. E. Eric Grissell

Systematic Entomology Laboratory, USDA

Smithsonian Institution

P.O. Box 37012

National Museum of Natural History CE 520, MRC168

Washington, DC 20013-7012

Phone: (202) 382-1781 Fax: (202) 786-9422 E-mail: egrissel@sel.barc.usda.gov

9088 01058 9604